‚ivist— st—li—n— di €—leontologi— e ƒtr—tigr—fi— volume IPH noF Q I plF ppF QVIERHV he™em˜er PHIR

„ri ve„i €visƒ„ygixi e€ixxsxi grewysƒ p‚yw q‚y„„e wy‚e ge†y‚ƒy

@ƒswf‚ sxs wy x„esxƒD gix„‚ev s„ev‰A

IB P I

viyxe‚hy ƒeve‚s D we‚sy pF ‚yvpy 8 ge‚wivy €i„‚yxsy

st

‚e™eivedX ƒeptem˜er I D PHIRY —™™eptedX xovem˜er PVD PHIR

uey wordsX v—te €leisto™eneD ‚upi™—pr—D qrott— wor— g—vorsoD of ƒouthern iurope —nd xe—r i—st @from QSë to SHë xA

st—lyF

@eul—gnier et —lF PHHVY rerrero et —lF PHHVAD ˜ut in the

v—te €leisto™ene its spre—d w—s mu™h widerD in™luding

e˜str—™tF „his p—per des™ri˜es — su˜EentireD p—rti—lly —rti™uE

lowE—ltitude regions —nd —lso ™o—st—l —re—s @w—sini IWVSY l—ted skeleton of ™h—mois found together with other verte˜r—te reE

m—ins in — silty lens with limestone ™l—sts ˜etween the IHR —nd IHS w—sini 8 vov—ri IWVVY w—sseti 8 ƒ—l—ri PHIP˜AF

ƒƒ of qrott— wor— g—vorso @v—tiumD st—lyAF „his silty lens is

„he genus is ™urrently divided into two spe™ies

™hronologi™—lly ™orrel—t—˜le to — temper—te os™ill—tion of wsƒ QF

@w—sini 8 vov—ri IWVVY qru˜˜ PHHSAX the elpine ™h—E

„he skull —nd postE™r—ni—l rem—ins from qrott— wor— g—vorso were

moisD ‚upi™—pr— rupi™—pr— @vinn—eusD IUSVAD divided ™omp—red with the fossil rem—ins of ™h—mois from numerous v—te

into seven region—l su˜spe™iesD spre—d from en—toli— €leisto™ene —nd older rolo™ene sites of st—ly —nd of €yrenees —nd

s˜eri—n €eninsul—F eddition—l ™omp—risons were m—de with l—rge s—mE —nd the g—u™—sus to the elpsY th—t of the €yreneesD

ples of ext—nt ™h—moisD ‚upi™—pr— pyren—i™— orn—t—D ‚F pyren—i™— pyrE

‚upi™—pr— pyren—i™— fon—p—rteD IVRSD widespre—d in

en—i™— —nd ‚F rupi™—pr— rupi™—pr—F worphologi™—l —nd morphometri™

the s˜eri—n €eninsul—D with its su˜spe™ies from the €yrE fe—tures of the skull —nd the hornE™oreD —s well —s other ™h—r—™ters of

enees —nd the g—nt—˜ri—n wount—insD whi™h h—s surE the met—™—rpus suggest to refer the fossil rem—ins from qrott— wor—

g—vorso to epennine ™h—moisD ‚F pyren—i™— orn—t—F „his is the e—rE vived up to present in st—ly only within the e˜ruzzi

liest —s™ert—ined re™ord in gentr—lEƒouthern st—ly of the orn—t—Elike

x—tion—l €—rk in the form of the su˜spe™ies ‚F pyreE

™h—moisD — su˜spe™ies so f—r known with ™ert—inty only in the roloE

n—i™— orn—t— xeum—nnD IVWW @see eul—gnier et —lF PHHVY

™eneD or possi˜ly in the v—tegl—™i—lF

rerrero et —lF PHHVAF ‚e™entlyD the epennine ™h—moisD

‚F pyren—i™— orn—t—D h—s ˜een su™™essfully reintrodu™ed

in other x—tion—l €—rks in gentr—l st—ly @w—ri 8 vov—ri

sntrodu™tion

PHHWAF

„he e—rliest fossil re™ords of ™h—mois in ‡estern „he v—te €leisto™ene ˜one rem—ins of ™h—mois

iuropeD referred to ‚upi™—pr— ™fF pyren—i™—D ™ome from from qrott— wor— g—vorso @tenneD v—tiumD st—lyAD more

l—yer «p» of the g—une de l9er—go @„—ut—velD €yreÂneÂesE spe™ifi™—lly — su˜EentireD p—rti—lly —rti™ul—ted skeletonD

yrient—lesD pr—n™eAD d—ted —˜out RRHDHHH ye—rs f€ —nd —re des™ri˜ed in this p—perF

™orrel—ted with the w—rine ssotope ƒt—ge IP @wsƒ IPA ‚em—ins of ‚upi™—pr— de fl—invilleD IVIT o™™ur in

@greÂgut IWUWY woigne et —lF PHHTY greÂgutEfonnoure ‡estern iurope sin™e the widdle €leisto™ene —nd —re

PHHUAF „he s—me t—xon w—s —lso re™orded from the site rel—tively ™ommon in the v—te €leisto™ene of st—ly @w—E

of yrgn—™ Q @erdeÁ™heD pr—n™eA in l—yers d—ted ˜etween sini 8 vov—ri IWVVY greÂgutEfonnoure PHHUY w—sseti 8

QQWDHHH —nd PWVDHHH ye—rs f€ @wsƒ WA @‚iv—ls PHHRAF ƒ—l—ri PHIP—D PHIP˜AF

„he finds then ˜e™ome slightly more numerous in the gurrent distri˜ution of this genus in™ludes sever—l

portions of the middle to highE—ltitude mount—in r—nges fin—l st—ges of widdle €leisto™ene @wsƒ TA —nd —mong

I B gorresponding —uthorF hip—rtimento di ƒ™ienze dell— „err—D ––ƒ—pienz—99 niversit—Á di ‚om—D €i—zz—le eldo woro SD sEHHIVS ‚om—D st—lyF

iEm—ilX leon—rdos—l—ridvirgilioFit

P hip—rtimento di ƒ™ienze ƒtori™heD filosofi™oEso™i—liD dei ˜eni ™ultur—li e del territorioD niversit—Á ––„or †erg—t—99D †i— golum˜i— ID sEHHITQ ‚om—D st—lyF

QVP ƒ—l—ri vFD ‚olfo wF pF 8 €etronio gF

pigF I E qrott— wor— g—vorsoX eA loE

™—tion of the siteY fA p—rti—l

longitudin—l se™tion @upA —nd

™omplete pl—n @downA of the

™—veF

these we ™—n mention other pren™h fossil rem—insX from —nd the w—ritime elpsD —s well —s in the xorthern

the qrotte du g—p de l— fielle @xestierD r—utesE€yreÂE epennines @w—sini IWVSY w—sini 8 vov—ri IWVVAD or

neÂesAD referred to ‚upi™—pr— ™fF pyren—i™—Y from the possi˜ly slightly f—rther southD —t le—st up to €is—no

qrotte des geÁdres @ve €l—n d9eupsD †—rA —nd the qrotte wountD „us™—nyD „yrrheni—n side @w—sseti 8 ƒ—l—ri

du v—z—ret @xi™eD elpesEw—ritimesAD —ttri˜uted to ‚F PHIP˜AF „he epennine ™h—mois w—s ™onfined to — porE

rupi™—pr—Y from the qrotte de l— xi™he @wontm—urinD tion of peninsul— ˜etween northern g—l—˜ri— —nd the

r—uteEq—ronnAD —ttri˜uted to ‚F pyren—i™— —nd from ƒi˜illini wount—ins @southern m˜ri—Ew—r™he epenE

‚om—inEl—E‚o™he @‚om—inD hou˜sAD referred to ‚upiE ninesA @w—sini IWVSY w—sini 8 vov—ri IWVVAD or pro˜E

™—pr— ™fF rupi™—pr— @see glot 8 w—rs—n IWVTY w—sini 8 —˜ly slightly f—rther northD up to the ™entr—l m˜ri—E

vov—ri IWVVY hefleur et —lF IWVWY „—voso et —lF IWWHY w—r™he epenninesD edri—ti™ side @w—sseti 8 ƒ—l—ri

greÂgutEfonnoure IWWPD PHHUY †—lensi PHHWY †er™outeÁre PHIP˜AF

8 queÂrin PHIHAF w—sini @IWVSD with referen™esA quoted QP st—li—n

possil rem—ins —ttri˜uted to ‚upi™—pr— in st—ly —re sites with fossil —nd su˜Efossil rem—ins of ™h—mois —nd

rel—tively numerous during the v—te €leisto™eneD espeE w—sseti 8 ƒ—l—ri @PHIP˜D with referen™esA RQ lo™—lities of

™i—lly from sites referred to the v—st ql—™i—l @‡urm gentr—lEƒouthern st—ly with the ™h—mois finds referred È

eu™torumAD ˜ut the t—xon pro˜—˜ly re—™hed the peninE to the v—te €leisto™ene —nd the older rolo™ene —™™ordE

sul— during the fin—l st—ges of the widdle €leisto™ene ing to the liter—ture —fter IWVVF ep—rt from these —˜oveD

@wsƒ TA @w—sini IWVSY €etronio et —lF PHHUD PHIIY €—E in st—ly we ™—n —dd some preE—lpine northEe—stern —nd

lom˜o et —lF PHIIAF „he —ssumed distri˜ution of ‚upiE viguri—n sites @eFgFD „—gli—™ozzo 8 g—ssoli IWWHD IWWPY

™—pr— in st—ly in the v—te €leisto™ene —nd the older g—ssoli 8 „—gli—™ozzo IWWID IWWR—D IWWR˜Y piore 8 „—E

rolo™eneD ˜—sed on the ™urrent distri˜ution of ™h—mois gli—™ozzo PHHSD with referen™esY ‡ierer 8 fos™—to PHHTY

popul—tions —nd the wellEpreserved rem—ins of only SEU †—lensi PHHWD with referen™esY qurioli et —lF PHIH—D

sitesD seems to ˜e —s followsF elpine ™h—mois w—s wideE PHIH˜Y fertolini et —lF PHIPA —nd the insul—r finds of il˜—

spre—d in the elpine —r™hD ˜etween the i—stern elps isl—nd @w—sseti in litterisAF sndeedD ex™ept for — few ™—sesD

v—te €leisto™ene epennine ™h—mois QVQ

ƒtr—tigr—phi™ notes of qrott— wor— g—vorso su™h —s qrott— di iquiD wonte ƒum˜r— —nd ‚ip— qr—nde

@w—sini IWVSA —nd ‚ip—ro di pum—ne @g—ssoli 8 „—gli—E

qrott— wor— g—vorso opens to UIS m —FsFlF ne—r

™ozzo IWWIY ‚om—ndini et —lF PHIRAD the rem—ins of ™h—E

tenne @v—tiumD gentr—l st—lyA —nd it le—ns out on the mois —re rel—tively poor —ndGor highly fr—gmentedF preE

right slope of the upper eniene ‚iver v—lley inside the quently they —re in su™h — st—te of preserv—tion th—t the

ƒim˜ruini wount—ins ‚egion—l x—tur—l €—rk @pigF IeAF l—™k of di—gnosti™ osteologi™—l m—teri—ls does not —llow

„his k—rst ™—vity hold — su™™ession of rooms —nd du™ts definite —ttri˜utions to one or the other spe™iesF gh—E

@pigF IfA —nd is ™—rved in the ––g—l™ilutiti e g—l™—reniti mois is —lso well represented from qrott— delle f—lze

del gon—™i—noEg—mp—ni—no99 form—tion @gWEIIGQVD with ˜one rem—ins of —lmost four young —nd two —dult

pper gret—™eousA @h—mi—ni et —lF IWWVY ‚olfo et —lF individu—ls ˜utD unfortun—telyD the hornE™ores with

PHHWAF pollowing the fortuitous dis™overy of hum—n front—l ˜ones mentioned in w—sini 8 vov—ri @IWVVA

˜ones inside the ™—veD —r™h—eologi™—l investig—tions —nd pu˜lished ˜y voreti 8 ƒ—lerno @IWVWA were not

st—rted in PHHT ˜y ––„or †erg—t—99 niversity of ‚omeD newfoundF „hereforeD the su˜Eentire skeleton from

in —greement with the ––ƒoprintendenz— per i feni erE qrott— wor— g—vorso is re—lly —n ex™eption—l —nd interE

™heologi™i99 of v—tium —nd with the ™oll—˜or—tion of esting ™—se studyF

pigF P E qrott— wor— g—vorsoX str—tiE

gr—phy of the room IF B—pE

proxim—te position of the

silty lens with the p—rti—lly

—rti™ul—ted skeleton of ™h—E moisF

QVR ƒ—l—ri vFD ‚olfo wF pF 8 €etronio gF

––ƒh—k— ulu99 speleologi™—l groupF „he inner rooms held ti™ul—rly wi™rotus @wi™rotusA —rv—lis —nd ervi™ol— —mE

the rem—ins of —t le—st PI individu—lsD the l—rgest xeoE phi˜ius with r—re „—lp— ™—e™—D „F rom—n—D ‚hinolophus

lithi™ hum—n ™ommunity of v—tium —nd one of the ri™hE ferrumequinumD wyotis ˜e™hsteiniiD wF em—rgin—tusD

est of iurope @‚olfo et —lF PHHWD PHIHD PHIP˜D PHIP™Y winiopterus s™hrei˜ersiiD wi™rotus @„erri™ol—A s—vii —nd

ƒ—l—ri et —lF PHIPAF sn the room ID just ˜elow the disE epodemus sylv—ti™usGfl—vi™ollis @ex™—v—tion PHHVY see

tur˜ed surf—™e —nd some rolo™ene l—yers @xeolithi™D ƒ—l—ri PHIRAF

fronze ege —nd histori™—l —geD see ‚olfo et —lF PHIHD felow the —forementioned ™ontexts is the IHR ƒ D

PHIP—D PHIQY ƒilvestri et —lF in pressAD the v—te €leisto™ene investig—ted ˜etween PHHV —nd PHIPF „his l—yer ™onsists

sequen™e w—s found @pigF PAF of in™oherent soilD m—inly ™h—r—™terized ˜y ™entimetri™

„he ƒtr—tigr—phi™ nits @ƒƒ A IHWD IIHD IPHD limestone ™l—stsF sn this ƒ D —˜out TH ™m thi™kD no huE

IQSD PSR —nd PSS —re ™h—r—™terized ˜y — ™omp—™t redE m—n —rtef—™tsD ˜ut very —˜und—nt mi™roverte˜r—te reE

dishE˜rown silty ground with limestone ™l—sts with m—ins together with sever—l ˜one fr—gments of m—™roE

sh—rp edges th—t v—ry in di—meter from few millimeters m—mm—l h—ve ˜een foundF €relimin—ry —n—lysis shows

to one ™entimeterD where m—ny fossil rem—ins of verteE th—t the domin—nt t—xon is wF @wFA —rv—lis @—lmost RVH

˜r—te were found together with — moder—te num˜er of m—ndi˜lesD —˜out RH7 of determined rem—ins —nd more

lithi™ —rtef—™ts refer—˜le to l—te pper €—l—eolithi™ @fin—l th—n UH7 of individu—ls —mong the m—mm—lsA followed

ipigr—vetti—nAF pront—l s™r—per on — retou™hed ˜l—de of ˜y sever—l rem—ins of eF —mphi˜ius —nd ˜irdsD vepus spF

j—spered flintD fr—gments of ˜—™ked ˜l—de letsD — fr—gment —nd w—rtes m—rtesD s™—r™e wF m—rmot—D eF sylv—ti™usG

of ˜—™ked point —nd — multiple ˜urin —re the findingsF sn fl—vi™ollis —nd gF i˜exD —nd r—re ‚eptili—D „F ™—e™—D „F

this ipigr—vetti—n l—yerD —˜out QH ™m thi™kD the most rom—n—D ƒorex s—mniti™usD wyodes gl—reolusD wustel—

—˜und—nt t—xon is w—rmot— m—rmot— @WR spe™imensD putoriusD ‚upi™—pr— spF —nd fos primigeniusF por str—tiE

—˜out h—lf of the determined rem—ins —nd one third of gr—phi™ position —nd for the ™omposition of m—mm—l

A —rv—lis —™™omp—nied the individu—lsA followed ˜y gervus el—phusD g—pr— —ssem˜l—geD domin—ted ˜y wF @wF

˜y ––™old99 m—mm—lsD su™h —s m—rmot —nd i˜exD this l—yer i˜exD ‚upi™—pr— spF —nd vepus spFD present with — sigE

w—s ™orrel—ted with the v—st ql—™i—l w—ximum @vqwA nifi™—nt num˜er of fr—gmentsD ƒus s™rof—D g—preolus ™—E

@yrl—™™hio PHIPY ‚i™™i PHIPAF preolus —nd †ulpes vulpes with few findsD —nd wustel—

IHS ƒ ˜elow followsY this is the deepest level niv—lis with only — single fr—gment of m—ndi˜le @digs

ex™—v—ted so f—rD me—suring —˜out PH ™m —t time @sumE PHHVEPHHWY see ƒ—l—ri et —lF PHIIAF e r—diometri™ d—ting

mer PHIQAF st is ™h—r—™terized ˜y — silty d—rk reddish soil performed on — m—rmot m—ndi˜le h—s provided the —ge

with r—re ™entimetri™ limestone ™l—sts —nd — l—rge of IQDRTH‹SH ye—rs f€ @gx‚ƒ vyon E RSTVD ™—li˜r—ted

—mount of fossil rem—ins of verte˜r—teD ˜ut without huE IRDIUS E IQDWIH fg IóAF „his —grees with the —ttri˜ution

m—n —rtef—™tsF €relimin—ry —n—lysis shows th—t the most of the lithi™ —rtef—™ts to the ™ultur—l horizon of the fin—l

—˜und—nt t—xon is vepus spF @SI spe™imensD —˜out PH7 ipigr—vetti—nD —nd of the m—mm—l —ssem˜l—ge to the

of determined rem—ins —nd IS7 of individu—ls —mong v—tegl—™i—l @see ƒ—l—ri et —lF PHIIAF „he finds of m—mm—ls

the m—mm—lsA followed ˜y gF i˜ex —nd ˜irdsD —nd then —˜ove —re —™™omp—nied ˜y sever—l ˜irds ˜onesD few —mE

˜y eF —mphi˜iusD gF el—phusD g—nis lupus —nd qlis glis phi˜ious ˜ones —nd sever—l mi™rom—mm—l rem—insD p—rE

pigF Q E qrott— wor— g—vorsoD room

IX the silty lens with limeE

stone ™l—sts of medium —nd

gre—t size —nd the p—rti—lly

—rti™ul—ted ˜one rem—ins of ™h—moisF

v—te €leisto™ene epennine ™h—mois QVS

sn the IWVHsD sever—l ™omp—risons of — morpholoE with some findsD s™—r™e wF @wFA —rv—lisD wF m—rmot—D

gi™—lD ˜iometri™—lD e™oEethologi™—lD ˜io™hemi™—lD —nd p—E ‚upi™—pr— spFD —nd r—re eF sylv—ti™usGfl—vi™ollisD ƒorex spF

leontologi™—l n—ture @vov—ri 8 ƒ™—l— IWVHY ƒ™—l— 8 voE —nd „F rom—n—F e r—diometri™ d—ting performed on —

v—ri IWVRY w—sini IWVSY x—s™etti et —lF IWVSY w—sini 8 skull fr—gment of wolf from this l—yer h—s provided —n

vov—ri IWVVAD en—˜led the re™ognition of two distin™t —ge higher of RQDSHH ye—rs f€ @fet— en—lyti™ sn™FD wi—E

spe™ies of ™h—mois in the prim—ry distri˜ution from mi E QTSQUSAF

the ‡estern €—le—r™ti™X ‚F rupi™—pr—D divided into v—rE fetween ƒƒ IHR —nd IHSD there w—s — silty lens

ious su˜spe™iesD —nd ‚F pyren—i™—D divided into three with limestone ™l—sts of medium —nd gre—t size @pigF QAF

su˜spe™iesD in™luding ‚F pyren—i™— orn—t—F purthermoreD rereD in — surf—™e of —˜out two sqmD more th—n WH

vov—ri 8 ƒ™—l— @IWVHA —nd w—sini 8 vov—ri @IWVVA in skelet—l elements of ™h—mois were foundD in™luding

p—rti™ul—rD reproposed in modern form — hypothesis th—t skullD p—rt of the sternumD —˜out twenty ri˜s —nd —s

h—d —lre—dy ˜een —dv—n™ed ˜y g—mer—no @IWIT—AF e™E m—ny verte˜r—eD —nd the lim˜ ˜onesF

™ording to su™h theoryD the popul—tions of ƒouthEwesE sn this silty lens with limestone ™l—sts —nd the su˜E

tern ™h—mois @€yrene—n —nd epennine ™h—moisAD who entire skeleton of ™h—moisD s™—ttered ˜one rem—ins of

were living sin™e the time of ‚iss gl—™i—tion in ‡estern two red deer —nd —t le—st three h—reD — su˜Eentire p—rE

iuropeD were sep—r—ted ˜y — new immigr—tion w—ve of ti—lly —rti™ul—ted skeleton of m—rmotD few rem—ins of

xorthEe—stern ™h—mois @elpine ™h—moisA during the firds —nd eF —mphi˜iusD —nd — m—ndi˜le of qF glis h—ve

‡urm gl—™i—tionF es we s—w in the ––sntrodu™tion99D in È ˜een —lso foundF por the o™™urren™e of gF el—phus —nd

the wsƒ IP —nd wsƒ W of ‡estern iurope there were qF glisD this —ssem˜l—ge is e™ologi™—lly ™loser to the

™h—mois rem—ins referred to ‚upi™—pr— ™fF pyren—i™—D m—mm—l —ssem˜l—ge of IHS ƒ th—n to IHR ƒ F gonE

˜ut in the wsƒ T there were —lre—dy two spe™iesD ‚F sidering th—t this silty lens with limestone ™l—sts is

rupi™—pr— —nd ‚F pyren—i™—F pl—™ed ˜etween the vqw @PH‹P k— ™—l f€D see entonioli

efter IWVVD the ™l—ssifi™—tion —˜ove h—s ˜een 8 †—i PHHRA —nd —t le—st RQDSHH ye—rs f€D these ˜one

widely —™™epted ˜y zoologists @eFgFD qru˜˜ IWWQD PHHSY rem—ins ™—n ˜e ™orrel—ted to — temper—te os™ill—tion of

emori et —lF IWWWY €edrotti 8 vov—ri IWWW—D IWWW˜Y euE wsƒ QF

l—gnier et —lF PHHVY rerrero et —lF PHHVY gorl—tti et —lF

PHIIAF snste—dD most p—leontologists —nd —r™h—eozooloE

gists h—ve kept on —ttri˜uting the osteologi™—l m—teri—ls „—xonomy of ‚upi™—pr— spe™ies

of ™h—mois from gentr—lEƒouthern st—ly to ‚F rupi™—pr—

„he system—ti™ of the genus ‚upi™—pr— is de˜—tedD or to ‚upi™—pr— spFD —nd those from €yrenees —nd s˜erE

p—rti™ul—rly the t—xonomi™ position of the epennine i—n €eninsul— to ‚F rupi™—pr— pyren—i™— or ‚F rupi™—pr—

™h—moisF @see w—sseti 8 ƒ—l—ri PHIP—D PHIP˜D with referen™es —nd

et the ˜eginning of the l—st ™enturyD essenti—lly on dis™ussion thereinAF

the ˜—sis of ™o—t ™olor p—tternD „rouess—rt @IWIHA re™ogE elso sever—l studies of mole™ul—r ˜iology h—ve

nized two spe™ies of ™h—moisD ‚F orn—t—D lo™—ted only in ™onfirmed the distin™tion ˜etween ‚F rupi™—pr— —nd ‚F

the e˜ruzzi mount—insD —nd ‚F rupi™—pr—D spre—d in pyren—i™— —nd the ™lose geneti™ —ffinity ˜etween the

other mount—in r—nges of ƒouthern iuropeD from the ™h—mois of e˜ruzzi —nd those of the €yrenees —nd s˜erE

€yrenees —nd g—nt—˜ri—n wount—ins to the elpsD g—rE i—n €eninsul— @see r—mmer et —lF IWWSY wu™™i et —lF

p—thi—nsD qree™e —nd g—u™—susF yn the ˜—sis of extern—l IWWVAF gonverselyD ‚odrõÂguez et —lF @PHHWD PHIHA ™l—im

—nd ™r—ni—l morphologyD willer @IWIPA —nd g—mer—no th—t postgl—™i—l re™oloniz—tion w—s —™™omp—nied ˜y hyE

@IWIT—D IWIT˜A divided the genus into the following speE ˜ridiz—tion ˜etween the pyren—i™— —nd rupi™—pr— ™l—desD

™iesX the epennine ™h—moisD ‚F orn—t—D the €yrene—n —nd ™on™lude th—t only one ‚upi™—pr— spe™ies exists

™h—moisD ‚F pyren—i™—D the elpine ™h—moisD ‚F rupi™—E divided into Q line—gesD ‡estern @pyren—i™— —nd p—rv—AD

pr—D divided into v—rious su˜spe™iesD —nd the sm—ll size orn—t— —nd ™—rtusi—n—A —nd i—stern @the other gentr—l @

g—nt—˜ri—n ™h—mois w—s ™onsidered — fourth spe™iesD rupi™—pr— su˜spe™iesAF snste—dD €eÂrez et —lF @PHHPD PHIQA

‚upi™—pr— p—rv— g—˜rer—D IWIHD ˜y willer @IWIPAD —nd —nd grest—nello et —lF @PHHWA hypothesized th—t the

— su˜spe™ies or v—riety of ‚F pyren—i™— ˜y g—mer—no epennine ™h—mois ™ould ˜e ™onsidered — third spe™iesF

@IWIT™AF snste—dD vydekker @IWIQAD then gouturier ‚e™entlyD on the ˜—sis of ˜iometri™—lD ethologi™—l

@IWQVA followed ˜y hol—n @IWTQA —nd fl—hout @IWUPAD —nd ˜iomole™ul—r ™onsider—tionsD qroves 8 qru˜˜

regrouped the ™h—mois into — single spe™iesD ‚F rupi™—E @PHIIA —nd qroves et —lF @PHIIA proposed the r—nk of

pr—D divided into sevenD then nine orD possi˜lyD ten reE full spe™ies for —lmost —ll the region—l su˜spe™ies of

gion—l su˜spe™iesD in™luding ‚F rupi™—pr— orn—t—F iven the genusD in™luding orn—t—D —nd —lso qippoliti @PHIQA

some p—leontologists re™ognized only one spe™iesD ‚F suggests this r—nk for the epennine ™h—moisF sn this

rupi™—pr— @see €r—t IWTTY uurteÂn IWTVAF p—per we —dhere to the ™urrent ™l—ssifi™—tion of the ™h—E

QVT ƒ—l—ri vFD ‚olfo wF pF 8 €etronio gF

skullsD or —t le—st for portion of skull with the hornE

™oresF fesides the insertion —nd in™lin—tion of the —˜ove

mentioned hornE™oresD ™ert—in —ddition—l di—gnosti™

™h—r—™ters h—ve ˜een identifiedD su™h —s the thi™kness

—nd the regul—rity or ™omplexity of v—rious sutures or

the o˜liter—tion of the ethmoid—l v—™uity @˜etween the

l—™hrym—lD n—s—lD front—l —nd m—xill—ry ˜onesA th—t is

fully ™losed in €yrene—n —nd epennine ™h—mois —nd

v—riously open in elpine ™h—mois —nd in the other ‚F

rupi™—pr— su˜spe™ies @g—mer—no IWIR˜D IWIT—D IWIT˜Y

gouturier IWQVY „os™hi IWTSY €r—t IWTTY eltun— IWUPY

vov—ri 8 ƒ™—l— IWVHY w—sini IWVSY w—sini 8 vov—ri

IWVVAF sn the verte˜r—te fossil depositsD howeverD int—™t

skulls —re very r—reD ˜ut ™r—ni—l fr—gments —re more

frequently foundD —nd in these ™—ses the morphometri™

di—gnosti™ ™riteri— rel—te to the dist—n™e @—t the ˜—se —nd

—pexA ˜etween the hornE™ores @eltun— IWUPY vov—ri 8

ƒ™—l— IWVHY w—sini IWVSAD while the length —nd the di—E

meters of e—™h hornE™ore —nd their r—tios —re not very

signifi™—nt @eltun— IWUPAD ˜ut useful for distinguishing

˜etween epennine —nd €yrene—n ™h—mois @gouturier

IWQVY ƒ™—l— 8 vov—ri IWVRY w—sini IWVSAF woreoverD pigF R E w—p of the m—in v—te €leisto™ene —nd older rolo™ene

the r—tio ˜etween the —nteroEposterior di—meter —nd

st—li—n sites for ™omp—risonsF €rX €r—destelY ƒoX ‚ip—ro ƒoE

the dist—n™e —t the ˜—se of the hornE™ores is — good m—nY „—X ‚ip—ro „—glienteY puX ‚ip—ro di pum—neY ƒfX

qrott— ƒ—n fern—rdinoY €dX €—dov—Y egX g—vern— delle method for distinguishing ˜etween elpineD epennine

erene g—ndideY iqX qrott— di iquiY ƒuX wonte ƒum˜r—Y —nd €yrene—n ™h—mois @w—sini IWVSAF gert—in distin™E

guX wonte gu™™oY f—X qrott— delle f—lzeY ‚uX qrott—

tive morphometri™ fe—tures ˜etween elpine —nd €yreE

‚utin—Y snX sng—r—noY €—X qrott— €—gli™™iY g—X qrott— dell—

ne—n ™h—mois h—ve ˜een identified for some postE™r—E

g—l—Y €oX ‚ip—ro del €oggioY „iX qrott— „in—Y ƒwX ƒ—ntu—rE

ni—l ˜onesD su™h —s the met—™—rpus @greÂgutEfonnoure io dell— w—donn—F ƒqu—reX elpine —r™ —nd xorthern

epennineY tri—ngleX gentr—lEƒouthern st—lyY rhom˜usD IWWPAF

wgX qrott— wor— g—vorsoF

mois popul—tions in two spe™iesD ‚F rupi™—pr— —nd ‚F w—teri—ls —nd methods

pyren—i™—F

possil m—teri—l from qrott— wor— g—vorso

„he elpine ™h—moisD ‚F rupi™—pr— rupi™—pr—D h—s

„he su˜Eentire skeleton of ™h—moisX

perpendi™ul—r hornsD the ˜ony ˜—ses of whi™h —re not PISUX p—rt of spl—n™hno™r—niumY PISU—X or˜it—l ˜onesY PISU˜X

hornE™ore with p—rt of front—l ˜onesY PISU™X portion of right m—xill—ry very exp—nded —nd —re sep—r—ted ˜y — rel—tively l—rge

I Q I Q

with w Ew Y PISUdX portion of left m—xill—ry with w Ew Y PPHQX left —re— of the front—l ˜onesY the thro—t p—t™h is little wide

Ew Y PPSH—X w Y m—ndi˜leY PPIVX right fr—gmented m—ndi˜le with € Q I P

—nd does not re—™h the ™hest —nd the winter ™o—t p—ttern

PPSHX w Y without num˜erX s Y PPVWX —tl—sY PUQIX —xisY PIWPD PIWP—D

Q P

h—s three sm—ll spotsF „he epennine ™h—moisD ‚F pyreE PIWV—D PPHSD PPHTD PPIVdD PPQTD PPSI™D PPSQD PPSUD PPSWD PPUVD PPVQD

n—i™— orn—t—D inste—dD h—s slightly ˜—™kEtilted hornsD PPVTD PPVVD PPVW— —nd PUQI˜X IU verte˜r—e —nd m—ny verte˜r— fr—gE

mentsY PPIH —nd PPQUX three fr—gments of sternumY PPHWD PPIH—D PPIPD whose ˜ony ˜—ses —re exp—nded —nd le—ve little sp—™e

PPIUD PPIVeD PPTI—D PPVHD PPVUD PPWHD PPWW —nd PQPHX PU fr—gments of

˜etween themY the thro—t p—t™h is wide —nd re—™hes the

ri˜sY PPVSX portion of right s™—pul—Y PPHRX left fr—gmented humerusY

™hest —nd the winter ™o—t p—ttern h—s five l—rge —nd one

PPHVX right fr—gmented humerusY PPTHEPPTIX fr—gmented left r—diusY

sm—ll spotsF „he €yrene—n ™h—moisD ‚F pyren—i™— pyreE PPWI—X dist—l p—rt of right r—diusY PPWIX proxim—l p—rt of right uln—Y

PPWV˜X right lun—teY PPRWX left met—™—rpusY PPQPX proxim—l p—rt of right n—i™— —nd ‚F pyren—i™— p—rv—D l—stlyD h—s slightly —nteE

met—™—rpusY PPSVX dist—l p—rt of right met—™—rpusY PPHUD PPTHD PPVRD riorlyEin™lined hornsD —nd the ˜ony ˜—ses of the horns

PPWIX —nterior first ph—l—ngesY without num˜erX some fr—gments of pelE

—re exp—nded —nd le—ve little sp—™e ˜etween themY the

visY PISTX fr—gmented left femurY PIWUX fr—gmented left ti˜i—Y PIWWX dist—l

thro—t p—t™h is wideD ˜ut — little less th—n in epennine

p—rt of right ti˜i—Y PPSSX right ™—l™—neusY PPTSX left ™—l™—neusY PPIV—X left

™h—moisD —nd the winter ™o—t p—ttern h—s five l—rge —nd t—lusY PPTRX right t—lusY PPSTX right ™entr—l t—rs—l ˜oneY PPHIX proxim—l

p—rt of right met—t—rsusY PPWTX proxim—l p—rt of left met—t—rsusY PPHP one sm—ll spots @g—mer—no IWIR—D IWIR˜D IWIT—D IWIT˜Y

—nd PPSIX posterior first ph—l—ngesY PIWVD PPHP—D PPHP˜D PPIV˜D PPSI— gouturier IWQVY „os™hi IWTSY vov—ri 8 fruno PHHQY „osi

—nd PPSV—X se™ond ph—l—ngesY PPIV™D PPRHD PPSI˜ —nd PPSV˜X third ph—E

8 €edrotti PHHQY q—r™õ—Eqonz—Âlez 8 rerrero PHHUAF

l—ngesF

„he osteologi™—l distin™tion ˜etween elpineD €yrE sn —ddition to the su˜Eentire skeleton des™ri˜ed ˜elowD the folE

ene—n —nd epennine ™h—mois is possi˜le for int—™t lowing rem—ins of ™h—mois —t wor— g—vorso h—ve ˜een foundX

v—te €leisto™ene epennine ™h—mois QVU

± ipigr—vetti—n l—yerX SQX left w Y IHHP right € Y without numE ‡e h—ve —lso ™—l™ul—ted the r—tios humerusGmet—™—rpus —nd I R

r—diusGmet—™—rpus following g—p—sso f—r˜—to et —lF @IWVSAF „he me—E € Y right s —nd w Y left s DsDs ˜erX — portion of left m—ndi˜le with € P Q P P I P Q

sures of lim˜ ˜ones of ‚F pyren—i™— p—rv— from er™eredillo et —lF @PHIIA —nd € Y right upper de™iduous ™heekEtoothY — fr—gment of lower mol—rY R

were used only for these r—tios —nd not for ™omp—risons on the —n—toE proxim—l p—rt of right uln—Y dist—l fr—gment of se™ond ph—l—nxY these

mi™—l elementsD ˜eing known the sm—ll size of this g—nt—˜ri—n wounE fossil rem—ins h—ve ˜een referred to — fully grown —nd — young indiviE

t—ins su˜spe™iesF du—l @see ƒ—l—ri et —lF PHIIAY

„he —ge of de—th of the spe™imen from qrott— wor— g—vorso

Y PQISX left t—lusY RIX posterE ± ƒ IHRX without num˜erX right € Q

w—s estim—ted through ™omp—rison with m—ndi˜les of ™h—mois of ior first ph—l—nxY

known —geD ™onsidering the —ge of eruptionsD ™h—ngeD —nd —ttrition of

—nd ± ƒ IHSX PTSTEPUQWX fr—gmented right m—ndi˜le with w I

the teethD —™™ording to the t—˜les —nd the o˜serv—tions reported in

Y PUVPX —rti™ul—r ™ondyle of met—podi—l ˜oneF w P

er™eredillo 8 hõÂez @PHHWAF ell the v—te €leisto™ene rem—ins of ™h—mois from qrott— wor—

g—vorso —re lo™—ted in the v—˜or—tory of €rehistory of the niversity

of ‚ome ––„or †erg—t—99F

„he ™h—mois from qrott— wor— g—vorso

gomp—rison s—mples

worphologi™—l —nd morphometri™ fe—tures of ™h—mois from „—phonomi™ o˜serv—tions

qrott— wor— g—vorso h—ve ˜een ™omp—red with v—te €leisto™ene —nd

sn summer PHIID —t the time of the —r™h—eologi™—l

rolo™ene ™h—mois rem—ins from sever—l sites of st—lyD €yrenees —nd

ex™—v—tion of the silty lens with limestone ™l—sts of medE s˜eri—n €eninsul— @„—˜F I —nd pigF RAF

ium —nd gre—t size ˜etween ƒƒ IHR —nd IHSD the gr—niometri™ d—t— of fossil ™h—mois €TQIID skull from ‚ip—ro

skullD the right m—ndi˜leD the pelvis —nd sever—l long di pum—neD —nd €IRHD right hornE™ore with front—l ˜one from the

p—l—eoveneti—n settlement of €—dov—D were t—ken ˜y the —uthorsF foth ˜ones were fr—gmentedY left s™—pul—D left uln—D right feE

spe™imens —re stored in the x—tion—l wuseum of €rehistory —nd ithE murD some ph—l—nges —nd most of the sm—ll ˜ones of

nogr—phy ––vuigi €igorini99 of ‚omeF „he me—surements of hornE™ores

™—rpus —nd t—rsus —re missingF ƒome verte˜r—eD right

from qrott— ‚utin— @ƒ—™™—Á PHHTAD qrott— di iqui @w—sini IWVSY glot

humerusD uln—D r—dius —nd lun—te were —rti™ul—tedD —s

8 w—rs—n IWVTAD wonte ƒum˜r— @w—sini IWVSAD eitz˜it—rte —nd rE

well —s ˜oth met—™—rp—l ˜ones —nd ph—l—ngesD right ti˜i—D ti—g— @eltun— IWUPA —nd v—rious ‡urmi—n sites from pren™h €yrenees È

t—lusD ™—l™—neus —nd ™entr—l t—rs—l ˜oneF „he other ˜ones @glot 8 w—rs—n IWVTA —re t—ken from liter—tureF „hese d—t— —re exE

posed in „—˜F PF were s™—tteredX left humerus —nd r—dius were very ™loseD

gr—niometri™ d—t— of the ext—nt elpineD €yrene—n —nd epennine

—s well —s left ti˜i— —nd met—t—rsusD ˜ut the left femur

™h—mois were t—ken ˜y g—mer—no @IWIR˜D IWIT—AD gouturier @IWQVA —nd

w—s found next to the skull —nd the m—ndi˜les ne—r the

w—sini @IWVSAY the young individu—ls —nd the spe™imens from h—uphineÂ

right met—™—rpusF ividentlyD the ™—r™—ss w—s p—rti—lly

—nd fosni—Ererzegovin— were not ™onsideredF „hese d—t— were inteE

dismem˜ered ˜y n—tur—l —gents @—nthropogeni™ input gr—ted with the me—surementsD t—ken ˜y —uthorsD of PI skulls of epenE

™—n ˜e ex™ludedA ˜efore ˜eing ˜uried ˜y mud —nd ™l—stsD nine ™h—mois from e˜ruzzi x—tion—l €—rk @spe™imens IID IRD PHD PID PTD

PWD PWD RQD SVD PIVD PPHD PPID SRUD IHWTD IISTD IPHQD IPHSD IPHVD IPISD presum—˜ly from overlying sinkholesF „hese two sinkE

IPIT —nd IPIUA housed —t the ƒ™ientifi™ ƒervi™e of ––inte eutonomo

holes h—ve ™ontrolled the sediments deposition in room

€—r™o x—zion—le d9e˜ruzzoD v—zio e wolise99 of €es™—sseroliF „hese

I —t le—st until to the fronze ege @‚olfo et —lF PHIQAF d—t— —re exposed in „—˜F QF

efter ™le—ning —nd w—shingD the fossil ˜ones —pE

„he sizes of the m—ndi˜le —nd the post™r—ni—l ˜ones of the

pe—r whitish or h—v—n—Etortoiseshell —nd —re very fr—gileF ™h—mois from wor— g—vorso h—ve ˜een —lso ™omp—red with the ™h—E

€relimin—ry —nd m—™ros™opi™ —n—lysis on the ˜one surE mois rem—ins from sever—l sites of st—lyD €yrenees —nd s˜eri—n €eninsuE

l—F por these ™omp—risonsD the osteometri™ me—surements h—ve ˜een in f—™es did not reve—l —ny tr—™es of hum—n —™tivity nor

p—rt t—ken from liter—ture —nd in p—rt dire™tly ˜y the —uthors @see „—˜F

signs of ™—rnivoresF ƒever—l ˜re—k—ges presum—˜ly

IAD in p—rti™ul—r the s—mples from qrott— delle f—lzeD housed —t the

™—used ˜y the weight of the sediments —nd the ™ontinE qu—ldo „—dino ƒpeleologi™—l qroup9s offi™eD from sng—r—noD housed

uous tr—mpling in ™onfined sp—™es or when digging were in the i—rth ƒ™ien™e hep—rtment of ––ƒ—pienz—99 niversity of ‚omeD

then re—ssem˜led in the l—˜or—toryF —nd from ‚ip—ro di pum—neD ‚ip—ro ƒom—n —nd qrott— ƒ—ntu—rio dell—

w—donn—D stored in the x—tion—l wuseum of €rehistory —nd ithnogr—E

phy ––vuigi €igorini99 of ‚ome @see „—˜F R —nd eppendix PAF

sn —dditionD we me—sured — m—ndi˜le @spe™imen €IQRGPA —nd the

hes™riptions —nd ™omp—risons

post™r—ni—l ˜ones of two elpine ™h—mois from qr—n €—r—diso x—tion—l

€—rk @spe™imens €SHT —nd without num˜erAD stored in the x—tion—l

ƒkull —nd m—ndi˜leF „he skull is fr—gmented —nd

wuseum of €rehistory —nd ithnogr—phy ––vuigi €igorini99 of ‚omeD

two inferior portions of m—xill—ries with the mol—rs

—nd IQ post™r—ni—l skeletons of epennine ™h—mois from e˜ruzzi x—E

were re™overed together with — p—rt of spl—n™hno™r—E tion—l €—rk @spe™imens PID PTD PUD QQD QSD RUD SVD IWID PHID PQVD PQWD IIHH

nium with n—s—l —nd preEm—xill—ry ˜ones —nd p—rt of —nd without num˜erA housed —t the ƒ™ientifi™ ƒervi™e of ––inte eutoE

the m—xill—ry —nd l—™hrym—l ˜onesD the right or˜it—l nomo €—r™o x—zion—le d9e˜ruzzoD v—zio e wolise99 of €es™—sseroli @see

eppendix QAF ˜ones —nd the right hornE™ore with p—rt of the right

—nd left front—l ˜ones @€lF ID figF IAF „he portion of

wethods

spl—n™hno™r—nium shows — fully ™losed ethmoid—l v—E

„he re™ent —nd fossil m—teri—l w—s me—sured —™™ording to the

™uityD —s in the €yrene—n —nd epennine ™h—moisF „he methodology of hries™h @IWUTA —ndD only for the met—™—rpusD —lso

hornE™ore shows ellipsoid—l se™tion —t the ˜—se —nd — —™™ording to greÂgutEfonnoure @IWWPA @see pigF SAF we—surements —re

slightly medioEl—ter—l ™ompression th—t evolve to — expressed in mmF „hese d—t— —re reported in t—˜les —nd di—gr—msF

QVV ƒ—l—ri vFD ‚olfo wF pF 8 €etronio gF

Site Geographic area Age References Gruta do Caldeirão Centro, Portugal Late Pleistocene Davis 2002 Gruta das Salemas Lisboa, Portugal Late Pleistocene Cardoso & Antunes 1989 Cueva del Nacimiento Andalucia, Spain Late Pleistocene Aférez Delgado et al. 1981 Alvarez et al. 1992; Pérez Lepido & Cueva Millan Castilla y Leon, Spain Late Pleistocene Cerdeño 1992 Rascaño Cantabria, Spain Late Pleistocene Altuna 1981 La Riera Cantabria, Spain Late Pleistocene Altuna 1986 Zatoya Navarra, Spain Late Pleistocene Altuna & Mariezkurrena 2001 Cueva de Abauntz Navarra, Spain Late Pleistocene Altuna et al. 2002 Cueva de SantimamiĖe Vizcaya, Spain Late Pleistocene Castaños 1984 Aitzbitarte Guipúzcoa, Spain Late Pleistocene Altuna 1972 Ermittia Guipúzcoa, Spain Late Pleistocene Altuna 1972 Lezetxiki Guipúzcoa, Spain Late Pleistocene Altuna 1972 Marizulo Guipúzcoa, Spain Late Pleistocene Altuna 1972 Urtiaga Guipúzcoa, Spain Late Pleistocene Altuna 1972 Cueva de Ekain Guipúzcoa, Spain Late Pleistocene Altuna & Mariezkurrena 1984 Urtao II Guipúzcoa, Spain Late Pleistocene Mariezkurrena 1989 Cueva de Erralla País Vasco, Spain Late Pleistocene Altuna & Mariezkurrena 1985 Labeko Koba País Vasco, Spain Late Pleistocene Altuna & Mariezkurrena 2000 Hautes-Pyrénées, Grotte du Cantet Late Pleistocene Clot 1984 France Hautes-Pyrénées, Grotte de Labastide Late Pleistocene Clot 1988 France various Würmian sites Pyrénées, France Late Pleistocene Clot & Marsan 1986 Grotte de Castel 2 Aude, France Late Pleistocene Pernaud et al. 2005 Late Pleistocene, early Tagliacozzo & Cassoli 1992 and this Riparo Soman Trentino Südtirol, Italy Holocene work Pradstel Trentino Südtirol, Italy early Holocene Boscato & Sala 1980 Grotta Maggiore San Veneto, Italy Late Pleistocene Cassoli & Tagliacozzo 1994b Bernardino Cassoli & Tagliacozzo 1991 and this Riparo di Fumane Veneto, Italy Late Pleistocene work Riparo Tagliente Veneto, Italy Late Pleistocene Capuzzi & Sala 1980 Tagliacozzo & Cassoli 1990 and this Padova Veneto, Italy Holocene work Arene Candide Liguria, Italy Late Pleistocene Cassoli & Tagliacozzo 1994a Grotta di Equi Tuscany, Italy Late Pleistocene Masini 1985; Clot & Marsan 1986 Monte Sumbra Tuscany, Italy Holocene Masini 1985 Monte Cucco Umbria, Italy Late Pleistocene Capasso Barbato et al. 1985 Grotta delle Balze Umbria, Italy early Holocene Loreti & Salerno 1989 and this work Grotta Rutina Abruzzi, Italy Late Pleistocene Saccà 2006 Benini et al. 1997; Boscato et al. Grotta della Cala Campania, Italy Late Pleistocene 1997 Grotta del Poggio Campania, Italy Late Pleistocene Sala 1979 Grotta Tina Campania, Italy Late Pleistocene Martini et al. 1974 Grotta Paglicci Apulia, Italy Late Pleistocene Boscato 1994 Ingarano Apulia, Italy Late Pleistocene Petronio et al. 1996 and this work Grotta Santuario della Late Pleistocene, early Calabria, Italy Fiore et al. 2004 and this work

Madonna Holocene „—˜F I E vist of ™omp—r—tive sitesD their —ge —nd geogr—phi™ —re—F

v—te €leisto™ene epennine ™h—mois QVW

Horn- MC Ru Eq Eq Eq Eq Su Su Su Su Pd Fu Fu IP IP IP IP FP FP FP FP core n. max min mea n. max min mea n. max min mea n. max min mea L. 128.0 20 104.0 57.0 78.6 86.5 120.0 79.9 4 90.0 77.0 83.3 17 112.0 67.0 92.7 APD 24.4 19.6 31 23.6 15.0 19.2 3 26.0 22.8 24.5 25.1 27.7 19.8 10 23.5 16.5 18.6 29 23.5 14.0 18.7 TD 22.7 28 20.2 12.1 16.6 3 22.2 20.0 20.7 22.9 26.0 18.0 9 16.5 14.3 16.3 28 23.0 12.4 17.5 BC 84.0 78.5 95.0 65.0 6 72.0 50.0 58.2 27 78.0 45.0 60.9

MD 10.4 12.6 13 19.0 12.0 14.9 3 19.0 14.5 16.8 15.6 10.2 2 11.0

„—˜F P E rornE™ore me—surements @mmA of fossil ™h—mois from some v—te €leisto™ene —nd older rolo™ene sites of gentr—lEƒouthern st—lyD

elpine —r™ —nd xorthern epennineD €yrenees —nd s˜eri—n €eninsul—F wgX qrott— wor— g—vorsoY ‚uX qrott— ‚utin— @ƒ—™™—Á PHHTAY iqX

qrott— di iqui @w—sini IWVSY glot 8 w—rs—n IWVTAY ƒuX wonte ƒum˜r— @w—sini IWVSAY €dX €—dov— @€IRH E this workAY puX ‚ip—ro di

pum—ne @€TQII E this workAY s€X s˜eri—n €eninsul— @eltun— IWUPAY p€X pren™h €yrenees @glot 8 w—rs—n IWVTAF

we—surement —˜˜revi—tionsX vFX lengthY e€hX —nteroEposterior di—meterY „hX tr—nsverse di—meterY fgX ˜—s—l ™ir™umferen™eY whX minimum dist—n™e —t the ˜—se of the hornE™oresY nFX num˜er of spe™imensY m—xX m—ximumY minX minimumY me—X me—nF

Horn-core n. max min mea s.d. n. max min mea s.d. n. max min mea s.d. R. r. rupicapra 1) 1) 1) 1) 1) 3) 3) 3) 3) 3) L. 142 124.0 62.0 87.4 13.33 APD 142 27.0 13.0 20.7 3.32 135 27.0 15.0 20.9 3.43 TD 142 26.0 12.0 18.4 3.41 139 26.0 12.0 18.4 3.56 MD 142 21.0 9.0 16.0 2.58 132 24.0 9.0 16.2 2.59 R. p. pyrenaica 1) 1) 1) 1) 1) 2) 2) 2) 2) 2) 3) 3) 3) 3) 3) L. 4 100.0 60.0 87.0 18.29 14 101.0 55.0 82.9 12.51 APD 4 22.0 15.0 18.5 2.89 14 21.0 12.5 17.0 2.07 18 22.0 12.5 17.4 2.29 TD 4 19.5 14.5 16.9 2.06 14 19.5 10.5 15.1 2.13 18 19.5 10.5 15.5 2.21 MD 4 10.0 8.0 9.0 0.82 14 17.0 10.5 13.4 1.53 18 15.0 8.0 12.3 2.36 R. p. ornata 1) 1) 1) 1) 1) 4) 4) 4) 4) 4) 3) 3) 3) 3) 3) L. 5 141.0 70.0 120.2 29.58 14 124.0 96.0 107.6 8.37 APD 5 29.0 19.0 25.4 4.62 19 28.0 20.2 23.9 2.67 32 29.0 19.8 24.8 3.32 TD 5 24.5 17.5 21.4 3.17 19 24.4 17.8 20.7 2.23 32 26.0 17.7 21.2 3.18

MD 5 14.0 10.0 11.9 1.60 19 16.0 10.6 13.4 1.79 21 15.8 9.5 12.7 1.93

„—˜F Q E rornE™ore me—surements @mmA of ™urrent popul—tions of ‚F rupi™—pr— rupi™—pr—D ‚F pyren—i™— pyren—i™— —nd ‚F pyren—i™— orn—t—F

h—t— fromX IA g—mer—no @IWIR˜D IWIT—AY PA gouturier @IWQVAY QA w—sini @IWVSAY RA this workF we—surement —˜˜revi—tionsX vFX lengthY

e€hX —nteroEposterior di—meterY „hX tr—nsverse di—meterY whX minimum dist—n™e —t the ˜—se of the hornE™oresY nFX num˜er of

spe™imensY m—xX m—ximumY minX minimumY me—X me—nY sFdFX st—nd—rd devi—tionF

su˜Etri—ngul—r se™tion with ™—ud—l ˜—se —t the —pexY the from qrott— ‚utin— @ƒ—™™—Á PHHTAD for th—t of sron —ge

medioEl—ter—l ™ompression is less signifi™—nt th—n in the from €—dov— —ndD presum—˜lyD for the spe™imens from

™urrent epennine ™h—moisF „he hornE™ore from wor— qrott— di iqui @see w—sini IWVSY w—sini 8 vov—ri IWVVAD

g—vorso is the longest —mong the fossil spe™imensD even dou˜ling the dist—n™e ˜etween the ˜—se of the right

th—n the gre—t spe™imen from pum—ne @„—˜F PAD —nd it is hornE™ore —nd the midpoint of the interEfront—l sutureF

longer th—n the living elpine —nd €yrene—n ™h—moisD „he r—tio ˜etween the —nteroEposterior di—meter —nd the

with dimensions slightly gre—ter th—n the me—n of the minimum dist—n™e —t the ˜—se of the hornE™ores shows

living epennine ™h—mois @„—˜F QAF „he di—meters —re th—t the spe™imen from wor— g—vorso is ™le—rly pl—™ed

™lose to the m—ximum v—lues of wonte ƒum˜r— —nd in the r—nge of the living epennine ™h—mois @pigF TAF

the pren™h €yreneesD slightly sm—ller th—n the spe™iE pper mol—rs —re without interlo˜—r lingu—l ™olE

mens from €—dov— —nd pum—ne —nd l—rger th—n the umnY on the o™™lus—l surf—™e they h—ve — sm—ll ™ir™le of

I P

other fossil spe™imens @„—˜F PAF „he two dimensions en—mel ˜etween the two m—in fossettesF w —nd w —re

I

—re gre—ter th—n in the modern €yrene—n ™h—moisD sm—lE ˜ilo˜ed —nd of simil—r sh—peD ˜ut w is slightly sm—llerD

Q

ler th—n the m—ximum v—lues of the ™urrent elpine —nd —nd it h—s the first lo˜e more lingu—lly proje™tingF w is

epennine ™h—moisD ˜ut ™lose to the me—n v—lues of the ˜ilo˜ed —nd it h—s — dist—l vesti˜ul—r stylus rel—tively

l—tter @„—˜F QAF developed —nd dist—lly ™onvexF „he length of the upper

„he minimum dist—n™e —t the ˜—se of the hornE mol—r row is slightly lower th—n th—t of the other st—li—n

™ores w—s ™—l™ul—tedD —s for the wousteri—n spe™imen fossil rem—insY it is ™lose to the me—n of the v—te €leisE

QWH ƒ—l—ri vFD ‚olfo wF pF 8 €etronio gF

GP Maxilla MC Ti Ti Cu SM SM Ru Fu Fu Fu So AC PIP PIP PIP PIP ANP ANP ANP ANP NP n. max min mea n. max min mea 22 38.0 41.6 42.0 40.4 39.1 5 43.0 38.0 40.5 17 39.2 35.4 36.7 Mandible 1 158.0 144.0 5 152.0 142.0 147.2 2 168.0 153.0 5 158.0 146.5 153.1 3 49.0 37.3 5 41.0 39.0 40.4 4 107.8 104.2 5 109.5 101.5 105.1 5 113.0 98.4 5 104.4 100.0 101.7 6 128.0 123.0 5 128.2 121.5 124.2 7 58.0 66.7 64.0 65.9 65.2 62.1 5 70.0 57.0 63.3 58.1 5 62.0 58.5 60.5 8 38.8 36.6 39.0 43.9 42.0 43.2 43.3 41.0 42.5 18 48.0 39.0 42.5 38.9 5 42.0 34.5 39.2 9 19.0 22.8 21.8 19.2 22.0 21.9 25.0 18.2 18 24.0 17.1 20.1 18.9 5 25.0 19.5 21.7 10a 17.8 19.0 20.3 19.4 20.5 10 19.0 17.0 17.8 17.3 4 17.8 16.0 16.9 10b 7.0 7.0 8.2 7.8 8.7 4 8.0 6.3 7.2 6.8 4 7.1 6.0 6.6 11 43.0 39.9 5 41.2 35.0 37.6 12 63.5 61.5 5 55.2 49.2 53.6 13 58.6 58.2 5 53.6 49.8 51.8 14 83.0 5 85.2 80.0 82.0 15a 28.8 33.8 30.4 33.1 33.9 35.0 1 32.0 28.3 5 30.2 21.2 26.4 15b 22.5 20.0 20.4 21.3 25.8 23.4 23.3 4 28.6 20.0 23.2 19.7 5 20.9 18.3 19.1

15c 17.8 15.4 18.7 18.9 15.8 19.4 18.4 5 19.4 16.6 17.7

„—˜F R E we—surements @mmA of upper mol—rs —nd m—ndi˜le of living ™h—mois —nd of rem—ins from some v—te €leisto™ene —nd e—rly rolo™ene

sites of gentr—lEƒouthern st—lyD elpine —r™D €yrenees —nd s˜eri—n €eninsul—F wgX qrott— wor— g—vorsoY „iX qrott— „in— @w—rtini et

—lF IWURAY guX wonte gu™™o @g—p—sso f—r˜—to et —lF IWVSAY ƒwX ƒ—ntu—rio dell— w—donn— @€IQTGP —nd €IQTGR E this workAY ‚uX qrott—

‚utin— @ƒ—™™—Á PHHTAY puX ‚ip—ro di pum—ne @g—ssoli 8 „—gli—™ozzo IWWIAY ƒoX ‚ip—ro ƒom—n @‚ƒVTH E this workAY egX g—vern— delle

erene g—ndide @g—ssoli 8 „—gli—™ozzo IWWR—AY €s€X €yrenees —nd s˜eri—n €eninsul—Y q€x€X ‚F rupi™—pr— rupi™—pr— from qr—n

€—r—diso x—tion—l €—rk @€IQRGPAY ex€X ‚F pyren—i™— orn—t— from e˜ruzzi x—tion—l €—rk @nnF PTD PWD IPHSD IPIS —nd without

num˜erAY nFX num˜er of spe™imensY m—xX m—ximumY minX minimumY me—X me—nF

we—surement —˜˜revi—tionsX w—xill—D PPX length of the mol—r rowY w—ndi˜leD IX length gonion ™—ud—le E infr—dent—leY PX length —˜or—l

Y RX length —˜or—l ˜order of ˜order of ™ondyle pro™ess E infr—dent—leY QX length gonion ™—ud—le E —˜or—l ˜order of —lveolus of w Q

E infr—dent—leY SX length gonion ™—ud—le E or—l ˜order of —lveolus of € Y TX length gonion ™—ud—le E the most —˜or—l —lveolus of w Q P

indent—tion of ment—l for—menY UX length of the ™heektooth rowY VX length of the mol—r rowY WX length of the premol—r rowY IH—X

Y IH˜X ˜re—dth of w Y IIX length of the di—stem—Y IPX height gonion ventr—le E highest point of ™ondyle pro™essY IQX height length of w Q Q

Y IS˜X height in gonion ventr—le E deepest point of m—ndi˜ul—r not™hY IRX height gonion ventr—le E ™oronionY IS—X height ˜ehind w Q

Y IS™X height in front of € F front of w I P

to™ene ™h—mois from €yrenees —nd s˜eri—n €eninsul— nine —nd premol—rsF „he right one is very fr—gmentedX —

D€ —nd w D —nd isol—ted w —nd w —nd to the higher v—lues of the ™urrent epennine ™h—E fr—gment with € Q R I P Q

were foundF vower teeth —re over—ll rounded —nd mois @„—˜F RAF

slightly hypsodontD the mol—rs —re without interlo˜—r „he left m—ndi˜le is —lmost int—™t @€lF ID figF PAD

vesti˜ul—r ™olumnF € h—s — mesioElingu—l not ˜ilo˜ed missing only the ™oronoid —pophysisD the in™isorsD ™—E Q

€ve„i I

‚upi™—pr— pyren—i™— orn—t— from qrott— wor— g—vorso @v—tiumD ™enE

tr—l st—lyAD v—te €leisto™eneF IA PISU˜ hornE™oreX —E l—ter—l viewD ˜E

front—l viewY PA PPHQ m—ndi˜le in l—˜i—l viewY QA PPVW —tl—s in dors—l

viewY RA PUQI —xis in l—ter—l viewY SA PPVS dist—l s™—pul— in l—ter—l viewY

TA PPHR humerus in ™r—ni—l viewY UA PPHV humerus in ™—ud—l viewY VA

PIWI proxim—l uln— in l—ter—l viewY WA PPTHEPPTI r—dius in dors—l viewY

IHA PPRW met—™—rpusX —E dors—l viewD ˜E p—lm—r viewY IIA PIST femur

in ™r—ni—l viewY IPA PIWU ti˜i— in ™r—ni—l viewY IQA PIWW dist—l ti˜i— in

™—ud—l viewY IRA PPIV— t—lus in dors—l viewY ISA PPTR t—lus in pl—nt—r

viewY ITA PPSS ™—l™—neus in medi—l viewY IUA PPST ™entr—l t—rs—l ˜one

in proxim—l viewY IVA PPHI proxim—l met—t—rsus in dors—l viewY IWA

PIWT proxim—l met—t—rsus in pl—nt—r viewY PHA PPHU —nterior first ph—E

l—nx in l—ter—l viewY PIA PPSI posterior first ph—l—nx in l—ter—l viewY

PPA PPIV˜ se™ond ph—l—nx in l—ter—l viewY PQA PPIV™ third ph—l—nx in

pigF S E we—surements of met—™—rpus @from greÂgutEfonnoure l—ter—l viewF

IWWPD redr—wnAF ƒ™—le ˜—rX S ™mF

v—te €leisto™ene epennine ™h—mois QWI

QWP ƒ—l—ri vFD ‚olfo wF pF 8 €etronio gF

etl—s —nd —xis h—ve the s—me proportions —s the

™urrent ™h—mois @€lF ID figs Q —nd RAY the —tl—sD howeverD

is l—rger —nd the —xis is ™lose to the gre—ter dimensions

of elpine —nd epennine ™h—moisY furthermoreD the —tl—s

is —lso l—rger th—n those from f—lzeD ƒ—ntu—rio dell—

w—donn— —nd ƒ—ntim—minÏ eD ˜ut sm—ller th—n th—t from

pum—neF

„he —rti™ul—r portion of the s™—pul— @€lF ID figF SA

deep — glenoid ™—vityD rel—tively n—rrow —nd elong—ted in

the —nteroEposterior dire™tionD the re—r edge is protrudE

ing —nd the ™or—™oid —pophysis is thi™k —nd protruding

downw—rd —nd forw—rdF „he dimensions —re ™lose to the

minimum v—lues of the ™urrent elpine —nd epennine

pigF T E ƒ™—ttergr—m of —nteroEposterior di—meter versus minimum

™h—mois —nd of —ll the fossils used in the ™omp—risonsF

dist—n™e —t the ˜—se of hornE™ores @mmA of the spe™imen

„he humerus @€lF ID figs T —nd UA is slenderY in the from qrott— wor— g—vorso ™omp—red with some v—te

dist—l endD the ole™r—non foss— h—s ogiv—l sh—peD the €leisto™ene —nd the living ™h—moisF ƒqu—reX —ttri˜uted to

‚F rupi™—pr—Y ™ir™leX —ttri˜uted to ‚F rF pyren—i™— or ‚upiE ridges th—t ˜order this —re lo™—ted more or less —t the

™—pr— ™fF pyren—i™—F „he me—surements of the rolo™ene ‚F

s—me levelY the tro™hle— with inner lip prominent —nd

pyren—i™— orn—t— from ‚ip— qr—nde —nd qrott— delle f—lze

fossette rel—tively deepF „he humerus of ™h—mois from

were not pu˜lished @see w—sini IWVSY w—sini 8 vov—ri

wor— g—vorso is slightly ™ompressed in the medioEl—tE IWVVY voreti 8 ƒ—lerno IWVWAF

er—l dire™tion —nd less ™ompressed in the —nteroEposteE

rior dire™tion th—t the ™urrent epennine ™h—moisF „he

reliefD — sh—llow dist—lEvesti˜ul—r groove —nd the side

proportions ˜etween the length —nd the dist—l epiphysis

with mesi—l lo˜e slightly edges diverging upw—rdsF € R —ppro—™h r—ther to those of ‚F rupi™—pr— spe™imens from

sm—ller th—n the posterior lo˜eD —nd th—t is ™omposed of qr—n €—r—diso x—tion—l €—rk @pigF UAF „he m—ximum

two denti™les sep—r—ted ˜y — fossetteF w —nd w —re I P length is less th—n the spe™imen from pum—neD ˜ut it

is slightly sm—llerD ˜ilo˜ed —nd of simil—r sh—peD ˜ut w I h—s the s—me size —s the living elpine ™h—moisD while

—nd it h—s the posterior lo˜e most prominent lingu—lly it is ˜igger th—n ˜oth the living epennine ™h—mois th—t

—nd the l—ter—l edges —re sh—rply diverging upw—rdsD the spe™imen from f—lze @the l—tter ˜eing ™lose to the

while in w they —re su˜Ep—r—llelsF w is trilo˜edD with P Q minimum v—lues of the modern epennine ™h—moisAF

the third lo˜e of redu™ed thi™kness —nd dist—lly ™onvexF „he size of the dist—l epiphysis —re in the r—nge of fossil

yver—llD the m—ndi˜le is simil—r in sh—peD ˜ut l—rger th—n spe™imens from gentr—lEƒouthern st—lyD they —re sm—ller

those of ™urrent ™h—mois @„—˜F RY see —lso g—mer—no th—n the spe™imen from pum—neD ™lose to those of modE

IWIR˜D IWIT—AF „he rounded posterior profile of the ern elpine ™h—mois —nd the minimum v—lues of epenE

m—ndi˜leD with the ™—ud—l gonion slightly protruding nine ™h—moisD —nd slightly ˜elow the —ver—ge of the v—te

˜—™kw—rdsD is ™ommon in the epennine ™h—moisD ˜ut €leisto™ene ™h—mois from €yrenees —nd s˜eri—n €eninE

not so frequent —mong the elpine ™h—mois @see g—mer—E sul—F

no IWIR˜D IWIT—AF roweverD the length @—lveol—rA of the worphology —nd size of the proxim—l uln— @€lF ID

lower ™heekEteeth row is ™lose to the spe™imen from figF VA f—ll within the v—ri—˜ility of the ™urrent ™h—mois

qr—n €—r—diso x—tion—l €—rkD to the minimum v—lues —nd of the fossil spe™imens used in the ™omp—risonY

of the ™urrent epennine ™h—mois —nd the v—te €leistoE howeverD the width of the ™oronoid pro™ess is rel—tively

™ene ™h—mois from €yrenees —nd s˜eri—n €eninsul—D ˜ut n—rrowF ‚—dius h—s rel—tively slender morphology @€lF ID

Ew length is less th—n the other fossil rem—insF „he w figF WAY the proxim—l end h—s the inside —rti™ul—r edge I Q

™lose to the spe™imen from qr—n €—r—diso x—tion—l slightly protruding —nd the posterior —rti™ul—r profile

€—rkD to the me—n of the ™urrent epennine ™h—moisD with the ™h—r—™teristi™ n—rrowD —ngul—r —nd sh—llow inE

to the minimum v—lues of the v—te €leisto™ene ™h—mois ™isureY in the dist—l endD the intern—l ™ondyle for —rti™uE

from €yrenees —nd s˜eri—n €eninsul— —nd to the size of l—tion with the s™—phoid is nonEedged ridges —nd the

the spe™imen from qrott— „in—D ˜ut lower th—n the intern—l ™ondyle for —rti™ul—tion with the lun—te is sh—lE

other fossil rem—ins ex—mined @„—˜F RAF lowF „he dimensions —re l—rger th—n those of the ext—nt

€ostE™r—ni—l˜onesF „he postE™r—ni—l ˜ones h—ve —ll epennine ™h—mois —nd slightly lower th—n those of the

the epiphyses fused —nd they h—ve the typi™—l fe—tures of spe™imens from qr—n €—r—diso x—tion—l €—rkF „he proE

g—prin—e —nd in p—rti™ul—r of the genus ‚upi™—pr—D portions ˜etween the length —nd the proxim—l —nd dist—l

˜eing mu™h more slenderD ˜oth in the epiphyses th—t epiphysesD howeverD —re the s—me of the ext—nt elpine

in the middle of the di—phisysD th—n to those of g—pr— —nd epennine ™h—mois —nd the e—rly rolo™ene s—mple

from f—lze @sm—ller th—n the minimum v—lues of living i˜ex —nd gF hir™usD ˜ut less slender th—n yvis —riesF

v—te €leisto™ene epennine ™h—mois QWQ

pigF U E ƒ™—ttergr—m of dist—l tr—nsverse di—meter versus m—ximum pigF W E ‚—tio di—gr—m of de™im—l log—rithmi™ differen™es ˜etween

length of humerus @mmA of the spe™imen from qrott— me—surements of the met—™—rpus of ™h—mois from qrott—

wor— g—vorso ™omp—red with some v—te €leisto™eneD wor— g—vorso @wgAD the me—n of T ‚F pyren—i™— orn—t—

e—rly rolo™ene —nd living ™h—moisF wgX qrott— wor— g—E individu—ls @‚poA —nd the me—n of V ‚F rupi™—pr— rupi™—pr—

vorsoY f—X qrott— delle f—lzeY puX ‚ip—ro di pum—neF individu—ls @‚rrA ™omp—red with the me—n of V ‚F pyren—iE

q€x€X living ‚F rupi™—pr— from qr—n €—r—diso x—tion—l ™— pyren—i™— individu—ls @HAF we—surement systemD sym˜olE

€—rkY ex€X m—ximumD minimum —nd me—n of living ‚F ogy @see pigF SA —nd d—t— of ‚F rupi™—pr— rupi™—pr— —nd ‚F

pyren—i™— orn—t— from e˜ruzzi x—tion—l €—rkF pyren—i™— pyren—i™— from greÂgutEfonnoure @IWWPAF

epennine ™h—moisAF ‡hile the v—te €leisto™ene ™h—mois from the €yrenees —nd s˜eri—n €eninsul—F „he proxim—l

epiphysis is less ™ompressed in the —nteroEposterior diE from €yrenees —nd s˜eri—n €eninsul— show — higher

re™tion th—n in the ™urrent ™h—moisX the proxim—l tr—nsE sturdiness in the middle of the di—physis —nd in pigF V

verse di—meter @fpA me—sure PRFH mm on —ver—geD the follow — different —lignmentD with little v—ri—tion in

proxim—l —nteroEposterior di—meter @hpA me—sure IVFT lengthD ˜ut ˜ig differen™es in the width of the dist—l endF

mm on —ver—geD so th—t the r—tio ˜etween the tr—nsverse „he met—™—rpus @€lF ID figF IHA is very slenderY the

di—meter —nd —nteroEposterior di—meter is IFPWD th—t of proxim—l epiphysis h—s rel—tively prominent dorsomeE

the spe™imens from qr—n €—r—diso x—tion—l €—rk is IFQI di—l tu˜erosity —nd the fossettes ˜etween the —rti™ul—r

—nd IFQRD —nd th—t of the s—mple from e˜ruzzi x—tion—l f—™ets whi™h —lso extends ˜elow the —rti™ul—r surf—™eF sts

€—rk is ˜etween IFQW —nd IFRTD with — me—n of IFRP @see length is gre—ter th—n the ™urrent epennine ™h—mois —nd

the v—lues in eppendix QAF of the spe™imens from qr—n €—r—diso x—tion—l €—rkD

‚eg—rding the met—™—rpusD greÂgutEfonnoure

˜ut sm—ller th—n th—t of the v—te €leisto™ene ™h—mois

@IWWPA h—s shown th—t in ‚F rupi™—pr— —nd ‚F pyren—i™—

of pr—n™e the proportions of the met—™—rp—l ˜ones reE

l—tive to the length @me—surements ID PD QA —nd their size

in the middle of the di—physis @me—surements IPD IQA —re

more or less the s—meF sn elpine ™h—moisD in the proxE

im—l endD the surf—™e for ™—rp—l ˜ones is wider @me—sureE

ments S to W —nd IIAD espe™i—lly the surf—™e for the os

m—gnumD —nd in the dist—l end @me—surements IR to PRA

the tu˜er™les of the dist—l di—physis —re more developed

th—n the tro™hle—F gonverselyD in the €yrene—n ™h—mois

the tu˜er™les —re less developed l—ter—lly @greÂgutEfonE

noure IWWPAF „he me—surements of elpine ™h—mois from

qr—n €—r—diso x—tion—l €—rk —re ™lose to the me—n v—E

lues provided ˜y greÂgutEfonnoure @IWWPA for ‚F rupiE

™—pr— of pr—n™e @„—˜F SAF „he epennine ™h—mois generE pigF V E ƒ™—ttergr—m of dist—l tr—nsverse di—meter versus m—ximum

—lly show intermedi—te v—lues ˜etween those of the elE length of r—dius @mmA of the spe™imen from qrott— wor—

g—vorso ™omp—red with some v—te €leisto™eneD e—rly roE pine —nd €yrene—n ™h—mois @„—˜F S —nd pigF WAF rowE

lo™ene —nd living ™h—moisF wgX qrott— wor— g—vorsoY f—X everD we m—y noti™e — gre—ter ro˜ustness in the middle

qrott— delle f—lzeY €s€X m—ximumD minimum —nd me—n of

of the di—physis @me—surements IPD IQA —ndD in the proxE

v—te €leisto™ene ‚F pyren—i™— from €yrenees —nd s˜eri—n

im—l end @me—surements S to W —nd IIAD — strong redu™E

€eninsul—F q€x€X living ‚F rupi™—pr— from qr—n €—r—diso

tion of the —rti™ul—r surf—™e for the os m—gnumY in the x—tion—l €—rkY ex€X m—ximumD minimum —nd me—n of

living ‚F pyren—i™— orn—t— from e˜ruzzi x—tion—l €—rkF dist—l end @me—surements IR to PRA the tu˜er™les of the

QWR ƒ—l—ri vFD ‚olfo wF pF 8 €etronio gF

GPNP GPNP R.r.r. R.r.r. R.r.r. R.p.p. R.p.p. R.p.p. R.p.o. R.p.o. R.p.o. SM Ba MC MC measure max min mea max min mea max min mea 1 149.0 146.0 164.8 130.1 146.4 145.2 129.2 139.5 145.0 135.5 142.4 153.0 2 137.6 134.7 151.0 118.0 135.2 134.1 117.6 128.2 131.0 122.5 129.2 142.5 3 143.6 140.6 157.7 128.1 141.1 140.6 123.9 135.4 139.5 131.4 137.6 149.0 4 24.5 24.1 29.0 22.9 24.9 24.4 21.6 22.7 25.5 23.0 24.4 24.1 23.9 5 24.0 23.5 28.0 22.4 24.3 24.5 20.8 22.1 23.8 22.5 23.2 23.1 22.9 6 10.1 9.8 11.8 9.1 10.2 10.1 8.4 9.2 10.3 9.4 10.0 10.8 10.7 7 16.1 15.7 18.6 14.7 16.2 15.1 12.8 14.1 14.4 12.6 13.5 13.9 13.8 8 14.4 14.8 17.2 13.4 14.5 14.5 12.8 13.7 15.2 13.5 14.6 14.5 14.4 9 19.1 19.7 22.8 17.8 19.3 18.5 16.0 17.5 20.0 18.4 19.4 19.4 19.4 10 18.3 18.4 20.3 16.6 18.0 17.1 15.6 16.4 17.7 16.5 17.1 18.8 18.5 11 16.7 16.9 19.4 15.1 16.5 15.8 13.7 15.0 16.7 15.2 15.7 16.6 16.5 12 15.3 14.6 19.5 12.9 15.5 15.6 13.9 14.8 16.0 15.2 15.7 15.7 16.1 13 11.8 11.0 14.1 10.1 11.7 12.2 10.2 11.1 11.8 11.2 11.6 11.0 12.0 14 29.0 27.9 32.8 24.9 28.7 27.4 25.2 26.4 30.5 28.0 29.3 29.4 27.7 29.2 29.1 15 28.6 27.0 32.5 24.3 27.8 27.0 24.6 25.8 28.6 26.8 27.6 27.7 27.7 29.1 28.8 16 13.8 13.1 15.1 11.3 13.4 13.5 12.0 12.5 13.8 8.0 12.6 12.6 12.8 13.7 13.5 17 11.6 11.6 14.3 9.9 12.0 11.9 9.3 10.9 11.8 8.0 10.8 11.2 11.0 11.3 11.2 18 15.1 16.5 19.0 15.0 17.0 16.9 15.4 16.2 18.4 16.0 17.4 16.4 17.2 16.8 16.8 19 17.0 16.1 18.6 14.5 16.6 16.5 14.5 16.6 18.8 17.0 17.8 17.8 16.6 17.3 17.2 20 13.0 14.0 16.3 12.1 14.4 14.2 12.5 13.3 15.8 14.0 15.1 14.6 14..2 14.2 14.4 21 16.8 17.8 19.8 15.7 17.5 16.7 15.1 15.9 17.0 15.8 16.6 16.7 15.9 17.3 17.3 22 10.8 11.6 13.2 9.8 11.4 10.9 9.6 10.3 10.9 10.2 10.6 10.8 10.4 10.2 10.3 23 17.4 17.4 19.6 15.7 17.1 16.4 15.0 15.8 17.2 16.0 16.7 17.4 16.1 17.2 16.9 24 10.5 11.1 12.3 9.4 10.9 10.5 9.2 9.8 11.1 10.2 10.6 10.7 10.2 10.1 10.1 2232-

specimen P506/93 WN P136/12 WN 2249

2258

„—˜F S E we—surements @mmA of met—™—rpus in re™ent elpineD epennine —nd €yrene—n ™h—mois —nd fossil rem—ins from some v—te €leisto™ene

—nd e—rly rolo™ene sites of gentr—lEƒouthern st—lyF ‚rrX m—ximumD minimum —nd me—n of spe™imens of V ‚F rupi™—pr— rupi™—pr—

@d—t— from greÂgutEfonnoure IWWPAY ‚ppX m—ximumD minimum —nd me—n of spe™imens of V ‚F pyren—i™— pyren—i™— @d—t— from

greÂgutEfonnoure IWWPAY ‚poX m—ximumD minimum —nd me—n of spe™imens of T ‚F pyren—i™— orn—t— from e˜ruzzi x—tion—l €—rkY

wgX qrott— wor— g—vorsoY f—X qrott— delle f—lzeY ƒwX ƒ—ntu—rio dell— w—donn—Y q€x€X elpine ™h—mois from qr—n €—r—diso

x—tion—l €—rkF

we—surement system —nd sym˜ology @see pigF SA from greÂgutEfonnoure @IWWPAF

dist—l di—physis —re more developed th—n the tro™hle— ‚eg—rding the femur @€lF ID figF IIAD the ne™k is

@more th—t in elpine ™h—moisD me—surements IRD ISA shortly elong—ted in the proxim—l epiphysisY in the dist—l

—nd —lso it is less ™ompressed in the —nteroEposterior end the lips of the tro™hle— —re —lmost simil—r —nd the

dire™tion @—s €yrene—n ™h—moisD me—surements ITD IUA inter™ondyl—r foss— is rel—tively wideF „he dimensions

—nd with the —rti™ul—r ™ondyles slightly less r—ised ˜ut —re ™lose to those of the spe™imens from qr—n €—r—diso

more sp—™ed @me—surements IVD IW —nd PHAF „he met—E x—tion—l €—rk —nd of the m—ximum v—lues of the ™h—E

™—rpus from wor— g—vorso is longer @me—surements ID mois from e˜ruzzi x—tion—l €—rkD ˜ut sm—ller th—n the

PD QA —nd more ro˜ust in the middle of the di—physis €leisto™ene spe™imen from pum—ne @pigF IHAF

@me—surements IPD IQA th—n the —ver—ge v—lues of ™urE „he ti˜i— —ppe—rs slender @€lF ID figs IP —nd IQAD

rent popul—tions of ™h—moisF „he other portions of the with dist—l epiphysis showing the f—™ets for the —rti™uE

™urve of this met—™—rpus follow the ™urve of the epenE l—tion with m—lleol—r ˜one prominent —nd sep—r—ted ˜y

nine ™h—moisD ˜ut there is — gener—lly minor ™ompresE — deep fissureY the —nterior fi˜ul—r f—™et is sm—llF sn the

sion in the —nteroEposterior dire™tion @me—surements IHD middle of the di—physis —nd in the dist—l end the dimenE

IID IQD ITD IUA —ndD in the dist—l endD — gre—ter redu™tion sions —re ™lose to those of the spe™imens from qr—n

of the extern—l ™ondyle ™—n ˜e noti™ed @me—surements €—r—diso x—tion—l €—rkD to the fossil rem—ins from €ogE

PQD PRA @„—˜F S —nd pigF WAF gio —nd wonte gu™™oD —nd to the —ver—ge of living

v—te €leisto™ene epennine ™h—mois QWS

—nd elpine ™h—moisF sn the ™—l™—neus @€lF ID figF ITAD in

dors—l viewD the —rti™ul—tion with m—lleol—r ˜one is elonE

g—ted —nd n—rrowD with the ™ondyle dist—lly limited ˜y —

tr—nsverse depression net —ndD ˜elow thisD followed ˜y —

™onvex portionF „he dimensions —re slightly gre—ter

th—n the ™urrent epennine ™h—mois —nd fossils from

wonte gu™™oD ™lose to those of the existing elpine ™h—E

moisD from sng—r—noD ƒ—ntu—rio dell— w—donn— —nd €r—E

destel —nd to the lower v—lues of the v—te €leisto™ene

™h—mois from €yrenees —nd s˜eri—n €eninsul—D ˜ut

mu™h sm—ller th—n the spe™imen from pum—neF gentr—l

t—rs—l ˜one with typi™—l morphology of g—prin—e @€lF ID

figF IUA —nd sizes th—t f—ll within the v—ri—˜ility of the

pigF IH E ƒ™—ttergr—m of dist—l tr—nsverse di—meter versus gre—t

™urrent ™h—mois —nd the fossil spe™imens for ™omp—riE

length from ™—put of femur @mmA of the spe™imen from

sonF

qrott— wor— g—vorso ™omp—red with some v—te €leisE

„he proxim—l portion of met—t—rsus @€lF ID figs IV to™ene —nd living ™h—moisF wgX qrott— wor— g—vorsoY

puX ‚ip—ro di pum—neF q€x€X living ‚F rupi™—pr— from —nd IWA shows the front of the m—in —rti™ul—r f—™ets

qr—n €—r—diso x—tion—l €—rkY ex€X m—ximumD miniE

™rossed ˜y — tr—nsverse depression rel—tively proE

mum —nd me—n of living ‚F pyren—i™— orn—t— from

noun™ed —nd the sm—ll f—™et for —rti™ul—tion with the

e˜ruzzi x—tion—l €—rkF

™entr—l t—rs—l ˜one little protrudingF „he tr—nsverse di—E

meter @fpA —re sm—ller th—n those of the spe™imen from

pum—neD —nd —re ™lose to those of other st—li—n fossil

rem—insD to the existing elpine —nd epennine ™h—mois

—nd to the minimum v—lues of the v—te €leisto™ene ™h—E

mois from €yrenees —nd s˜eri—n €eninsul—F „he proxiE

m—l tr—nsverse di—meter @fpA me—sure PPFUS mm on

—ver—geD the proxim—l —nteroEposterior di—meter @hpA

me—sure PHFV mm on —ver—geD so th—t the r—tio ˜etween

the tr—nsverse di—meter —nd —nteroEposterior di—meter is

IFHW —nd f—lls within the v—ri—˜ility of st—li—n re™ent —nd

fossil ™h—moisX qr—n €—r—diso x—tion—l €—rk IFHU —nd

IFHWD e˜ruzzi x—tion—l €—rk ˜etween IFHQ e IFIV with —

me—n of IFIID f—lze IFHP —nd IFHQD pum—ne IFHS —nd

IFHUD wonte gu™™o IFIQ @see the v—lues in eppendi™es pigF II E ƒ™—ttergr—m of —nteroEposterior di—meter versus tr—nsE

verse di—meter of ti˜i— dist—l epiphysis @mmA of the spe™iE P —nd QAF

men from qrott— wor— g—vorso ™omp—red with some

„he proxim—l ph—l—nges @€lF ID figs PH —nd PIA —re

v—te €leisto™eneD e—rly rolo™ene —nd living ™h—moisF

rel—tively elong—ted —nd slenderD slightly ™ompressed in

wgX qrott— wor— g—vorsoY €oX qrott— del €oggioY guX

the medi—lEl—ter—l dire™tionY the —nterior ones —re ™omE wonte gu™™oY f—X qrott— delle f—lzeY €rX €r—destelY puX

‚ip—ro di pum—neY ƒoX ‚ip—ro ƒom—nY €s€X m—ximumD p—™t —nd str—ightD the posteriors slightly ™urved upw—rds

minimum —nd me—n of v—te €leisto™ene ‚F pyren—i™— —nd with rel—tively n—rrow dist—l epiphysisF „he dimenE

from €yrenees —nd s˜eri—n €eninsul—F q€x€X living ‚F

sionsD ˜oth of the —nteriors —nd the posteriorsD —re l—rger

rupi™—pr— from qr—n €—r—diso x—tion—l €—rkY ex€X

th—n those of ™urrent epennine ™h—mois —nd of most

m—ximumD minimum —nd me—n of living ‚F pyren—i™—

st—li—n ™omp—r—tive sitesD ™lose to the m—ximum v—lues orn—t— from e˜ruzzi x—tion—l €—rkF

of the v—te €leisto™ene ™h—mois from €yrenees —nd

s˜eri—n €eninsul—D ˜ut sm—ller th—n the spe™imen from

pum—neF sn —dditionD the size of the —nterior ph—l—nges epennine ™h—mois —nd v—te €leisto™ene ™h—mois from

—re ™lose to the spe™imens from qr—n €—r—diso x—tion—l €yrenees —nd s˜eri—n €eninsul—D ˜ut sm—ller th—n ƒom—n

€—rk whi™h h—veD howeverD the posterior ph—l—nges of —nd pum—ne —nd l—rger th—n €r—destel —nd f—lze @see

sm—ller size @„—˜F T —nd eppendi™esAF pigF IIAF

„he se™ond ph—l—nges @€lF ID figF PPA —re rel—tively „he t—lus @€lF ID figs IR —nd ISA is wide —nd rel—E

slender —nd slightly ™ompressed in the medi—lEl—ter—l tively lowD with the lips of the tro™hle— proxim—l div—riE

dire™tionF „he dimensions —re ™lose to spe™imens from ™—ted —nd with the side edges diverging upw—rdsY the

qr—n €—r—diso x—tion—l €—rkD to the fossil rem—ins of posteriorEdist—l fossette is sh—llowF „he dimensions f—ll

the ƒ—ntu—rio dell— w—donn— —nd the m—ximum v—lues within the v—ri—˜ility of the fossils used in the ™omp—rE

of the v—te €leisto™ene ™h—mois from €yrenees —nd isonD ˜eing slightly gre—ter th—n the ™urrent epennine

QWT ƒ—l—ri vFD ‚olfo wF pF 8 €etronio gF

Po Po Po Po Cu Ca Ca Ca Ca Ca Ca Fu Fu Fu Fu Fu SB SB SB PIP PIP PIP PIP Ph. I n. max min mea GL 50.6 50.0 46.0 46.7 46.0 59.8 58.2 57.7 54.2 55.6 54.5 54.9 52.7 54 50.8 36.5 45.5 Bp 13.5 14.8 14.9 13.3 12.2 12.1 12.4 14.6 16.9 17.3 16.8 16.5 15.1 16.7 15.3 16.4 52 15.7 11.3 13.7 SD 9.7 10.0 10.3 12.7 12.2 11.5 12.0 11.1 12.2 10.9 12.3 49 11.5 8.4 9.7 Bd 13.0 13.0 12.6 13.5 14.0 14.3 11.6 15.9 15.7 15.9 14.6 13.6 16.0 14.5 15.1 46 14.2 10.5 12.7 Ph. II GL 28.0 29.0 34.5 30.3 28.0 40.1 37.9 34.6 35.7 37.1 31.5 33.9 42 33.1 20.5 29.6 Bp 12.5 12.5 14.0 13.4 12.7 12.8 14.0 13.9 16.2 16.0 15.9 16.1 14.7 25.5 14.6 40 15.5 9.4 13.0 SD 8.6 7.6 9.0 7.6 10.4 10.3 9.7 10.2 9.6 9.6 9.2 13 9.5 6.6 8.2 Bd 9.5 9.5 10.8 9.0 10.1 11.0 12.1 12.6 11.6 11.7 11.5 11.1 10.5 16 11.2 7.6 10.1 Ph. III DLS 38.3 34.7 12 38.626.5 33.2 Ld 32.3 28.8 7 31.5 22.0 26.2

MBS 7.8 3 11.3 8.3 9.5

„—˜F T E we—surements @mmA of firstD se™ond —nd third ph—l—nges of some v—te €leisto™ene sites of ™entr—lEsouthern st—lyD elpine —r™ —nd

€yrenees —nd s˜eri—n €eninsul—D t—ken ˜y liter—tureF €oX qrott— del €oggio @ƒ—l— IWUWAY guX wonte gu™™o @g—p—sso f—r˜—to et —lF

IWVSAY g—X qrott— dell— g—l— @fenini et —lF IWWUY fos™—to et —lF IWWUAY puX ‚ip—ro di pum—ne @g—ssoli 8 „—gli—™ozzo IWWIAY ƒfX qrott—

w—ggiore ƒ—n fern—rdino @g—ssoli 8 „—gli—™ozzo IWWR˜AY €s€X €yrenees —nd s˜eri—n €eninsul— @see ––w—teri—ls —nd methods99AY €hF sX

first ph—l—nxY €hF ssX se™ond ph—l—nxY €hF sssX third ph—l—nxY nFX num˜er of spe™imensY m—xX m—ximumY minX minimumY me—X me—nF por

the me—surements of ph—l—nges from qrott— wor— g—vorso —nd other v—te €leisto™eneD e—rly rolo™ene —nd living ™h—mois t—ken ˜y

—uthorsD see the eppendi™esF

we—surement —˜˜revi—tionsX qvX gre—test lengthY fpX ˜re—dth of proxim—l endY ƒhX sm—llest ˜re—dth of the di—physisY fdX ˜re—dth of

dist—l endY hvƒX di—gon—l length of the soleY vdX length of dors—l surf—™eY wfƒX ˜re—dth in the middle of the soleF

s˜eri—n €eninsul—D they —re sm—ller th—n the l—rgest speE Horn-core males females

™imen from pum—neD ˜ut gener—lly l—rger th—n the other max min mea max min mea

st—li—n sites —nd the existing ™h—mois from e˜ruzzi x—E R. r. rupicapra

tion—l €—rk @„—˜F T —nd eppendi™esAF L. 124.0 62.0 94.8 102.0 62.0 77.8

„he third ph—l—nges @€lF ID figF PQA —re rel—tively TD 26.0 16.0 20.7 25.0 12.0 15.5

high —nd ™ompressed in medi—lEl—ter—l dire™tionD —nd R. p. pyrenaica

show the front p—rt of the lower f—™e of ogiv—l sh—peF L. 101.0 95.0 98.0 93.0 55.0 77.6

„he length is signifi™—ntly l—rger th—n —ll the existing TD 19.5 15.5 17.3 17.0 10.5 17.3

—nd fossil spe™imens used in the ™omp—risonD ˜ut the R. p. ornata

midpoint ˜re—dth of the sole is simil—r to the st—li—n L. 141.0 105.0 122.6 117.0 96.0 107.3

™h—mois —nd slimmer th—n the v—te €leisto™ene ™h—mois TD 24.5 17.8 22.4 20.3 18.3 18.9

of €yrenees —nd s˜eri—n €eninsul— @„—˜F T —nd eppenE

„—˜F U E rornE™ore me—surements @mmA of modern m—les —nd feE di™esAF

m—les of ‚F rupi™—pr— rupi™—pr— @d—t— from g—mer—no

IWIR˜AD ‚F pyren—i™— pyren—i™— @d—t— from gouturier

ƒex —nd —ge of the spe™imen

IWQVA —nd ‚F pyren—i™— orn—t— @d—t— from g—mer—no

ƒexF sn the ™h—mois ˜oth sexes h—ve hookEsh—ped IWIT— —nd this workAY m—xX m—ximumY minX minimumY

me—X me—nY vFX lengthY „hX tr—nsverse di—meterF hornsD those of the m—les ˜eing longer —nd thi™ker —nd

showing more ™losed —pi™—l hook @g—mer—no IWIR—Y

gouturier IWQVY vov—ri 8 fruno PHHQY „osi 8 €edrotti

er—no IWIR˜Y gouturier IWQVY „osi 8 €edrotti PHHQY PHHQY q—r™õ—Eqonz—Âlez 8 rerrero PHHUAF „—˜le U shows

er™eredillo 8 hiez PHHWAD —t W ye—rs the lower the v—ri—˜ility of the length —nd the tr—nsverse di—meter

™heekEteeth shown — little we—r —nd —t IS ye—rs — gre—t —t the ˜—se of hornE™ore ˜etween m—les —nd fem—les of

st—te of we—r ex™ept w @gouturier IWQVY er™eredillo ext—nt elpineD epennine —nd €yrene—n ™h—moisF „he I

8 hiez PHHWAF „he m—ndi˜le of our ™h—mois h—s medE d—t— of epennine ™h—mois —re inferred ˜y using ˜oth

iumEworn mol—rsD —nd the height of the unworn ™rown the v—lues reported in g—mer—no @IWIT—A —nd the v—lues

is ITFU mmY the lower mol—rs show intermedi—te of w of our s—mple of ‚F pyren—i™— orn—t— from e˜ruzzi P

we—r st—ge ˜etween the m—ndi˜les q @W ye—rsA —nd r x—tion—l €—rk @ex™ept for young individu—ls —nd those

@IS ye—rsA illustr—ted ˜y er™eredillo 8 hiez @PHHWD figF of ––undetermined sex99AF e™™ording to the l—rge size of

QAD —nd slightly gre—ter th—n the we—r st—ge of lower its hornE™ore @lengthX IPVFHY tr—nsverse di—meter —t the

™heekEteeth of epennine ™h—mois from e˜ruzzi x—E ˜—seX PPFUY see „—˜F PA the spe™imen from qrott— wor—

tion—l €—rk of IP ye—rs @spe™imen RUAF „he we—r st—ge g—vorso is very likely to ˜e — m—leF

of upper mol—rs —re simil—r to th—t of the spe™imens II egeF „he elpine —nd €yrene—n ™h—mois h—ve the

™omplete perm—nent dentition —t —˜out S ye—rs @g—mE —nd IRD respe™tively of IP —nd IR ye—rsF „husD the

v—te €leisto™ene epennine ™h—mois QWU

dent—l we—r indi™—tes —n —ge pro˜—˜ly slightly gre—ter h—ve sm—ller —nd n—rrower third ph—l—nges th—n the

ki—ng —nd the €rzew—lski9s horses whi™h run most of th—n IP ye—rsY —lso the m—ximum height of the horizonE

the ye—r on rel—tively soft groundD s—ndy or snowy @iiE D r—ther th—n posterE t—l ˜r—n™h —t the se™ond lo˜e of w Q

senm—nn IWVRY gonti et —lF PHIHAF sn the presen™e of the iorlyD indi™—tes —n —dv—n™ed —geF

s—meD or simil—rD environment—l ™onditions @w—lk —nd

run for p—rt of the ye—r on rel—tively soft or snowy

groundAD it seems th—t the v—te €leisto™ene €yrene—n ‚em—rks

™h—mois h—d — different —d—ptive responseD in order to

worphologi™—l —nd morphometri™ fe—tures of in™re—se the supporting surf—™eX they h—ve the third ph—E

skull —nd hornE™ore of ™h—mois from qrott— wor— g—E l—nx rel—tively short —nd — gre—ter —mplitude of the soleF

vorso —re typi™—l of the ext—nt epennine ™h—moisD in snste—dD the v—te €leisto™ene ™h—mois of ƒ—ntu—rio dell—

p—rti™ul—r the fully ™losed ethmoid—l v—™uity —nd the w—donn— —nd qrott— dell— g—l—D lo™—lities ne—r the se—

r—tio ˜etween the —nteroEposterior di—meter —nd the in ƒouthern st—lyD h—d the third ph—l—nges of the s—me

minimum dist—n™e —t the ˜—se of the hornE™oresF size of the modern ™h—moisD —s simil—r ™onditions —t

‚eg—rding the postE™r—ni—l skeletonD the ˜ones of high —ltitudes in the mount—ins ™ould ˜e found in ro™ky

this spe™imen @—s well —s most other v—te €leisto™ene ™o—st—l —re—sF

spe™imensA —re gener—lly gre—ter th—n the postE™r—ni—l ‚eg—rding the proportions of lim˜ ˜onesD g—p—sE

˜ones of e—rly rolo™ene —nd living ™h—moisD ˜ut it so f—r˜—to et —lF @IWVSD t—˜F IA h—ve shown th—t in the

w—s —lre—dy noti™ed th—t from the v—te €leisto™ene to ext—nt ‚F rupi™—pr— @sensu ˜efore IWVVA the r—tios huE

the rolo™ene the ˜ody size of some spe™iesD su™h —s merusGmet—™—rpus —nd r—diusGmet—™—rpus —re on —verE

g—pr— i˜ex @f—rtolomei 8 ƒ—l— IWUPAD iquus ferus @iiE —ge equ—l to IFPV —nd to IFPWD while for ex—mple in gF

senm—nn IWWIY gonti et —lF PHIHD —nd referen™es thereinAD i˜ex these r—tios —re —n —ver—ge respe™tively of IFSR —nd

†ulpes vulpes @€etronio et —lF PHHTA —nd fos primigenius IFPPD reve—ling th—t the l—tter spe™ies h—s the met—™—rpus

@€—ndolfi et —lF PHIID with referen™esA diminishedD p—rtiE signifi™—ntly shorter th—n the humerus —nd slightly

™ul—rly in the dimensions of the lim˜sF shorter th—n the r—diusF „hese r—tios in ™h—mois from

„he lim˜ ˜ones of the spe™imen from wor— g—E wor— g—vorso —re respe™tively IFPTI —nd IFPSSF ‡e h—ve

vorso —re mostly simil—r to the ™orresponding skelet—l —s™ert—ined th—t the v—lues given ˜y g—p—sso f—r˜—to et

elements of the ext—nt epennine ™h—moisD ex™ept for the —lF @IWVSA —re intermedi—te ˜etween the me—n v—lues of

humerus —nd the dist—l ti˜i—D th—t morphometri™—lly —re our s—mples of the ext—nt elpine —nd epennine ™h—E

™loser to those of elpine ™h—moisF „he eldest differen™e moisF „—˜le V shows the r—tios humerusGmet—™—rpus

from the fossil —nd modern ™h—mois is the length of the —nd r—diusGmet—™—rpus for the spe™imen from wor— g—E

ph—l—ngesD p—rti™ul—rly of the third ph—l—nxF sn the ™urE vorso —nd the s—mples for ™omp—risonF „he v—lues of

rent ™h—moisD the third ph—l—nges —re rel—tively sm—llD lengths of these skelet—l elements of wor— g—vorso ™h—E

short —nd thinD —nd so f—vor the motionD jumping —nd mois —re in eppendix ID those of the elpine ™h—mois

running on the ro™ksF „he stru™ture of the hooves gives from qr—n €—r—diso x—tion—l €—rk —nd epennine ™h—E

it gre—ter —˜ility to move on the mount—in groundX the mois from e˜ruzzi x—tion—l €—rk —re in eppendix QF sn

outside edge is h—rd —nd sh—rp —llowing the support the m—les of ext—nt ‚F pyren—i™— p—rv— the length of

even on sm—ll ro™ky out™ropsD the front pointing down humerus r—nges from ITQFS to IUSFW mmD with — me—n

slightly f—voring w—lking on i™eD the sole of the foot is of ITVFV mmD the length of r—dius from ITSFH to IVSFW

el—sti™D strong —nd slightly ™on™—veD in™re—sing —dherE mmD with — me—n of IURFP mmD —nd the length of meE

en™e to the steep ro™ky slopesY moreoverD the presen™e t—™—rpus from IQTFV to ISPFQ mmD with — me—n of IRQFH

of —n interdigit—l skin foldD in™re—sing the ˜e—ring surE mm @er™eredillo et —lF PHIIAD while in the v—te €leistoE

f—™eD f—™ilit—tes w—lking on soft —nd snowy soils @gouE ™ene spe™imens from €yrenees —nd s˜eri—n €eninsul—

turier IWQVY vov—ri 8 fruno PHHQY „osi 8 €edrotti the length of r—dius r—nges from IVUFS to IVWFH mmD

PHHQAF sn the spe™imen of wor— g—vorsoD the rel—tively with — me—n of IVVFQ mmD the length of met—™—rpus

long posterior first ph—l—nges —nd the long —nd thin from ISUFQ to ITPFS mmD with — me—n of ISWFU mm

third ph—l—nges m—y ˜e —n —d—pt—tion to motion on soft @see eltun— IWUPD IWVSY w—riezkurren— IWVWY h—vis

soils @gr—ssl—nds —nd forests of the more or less snowy PHHPAF

v—lley floorA ˜eing the ro™ky —re—s of high —ltitude of elthough the spe™imens of modern elpine ™h—E

the gentr—l epennines ™overed with snow —nd i™e durE mois —nd the v—te €leisto™ene ™h—mois from €yrenees

ing the v—st ql—™i—lF sn p—rti™ul—rD e—™h p—ir of longer —nd s˜eri—n €eninsul— —re not numerousD it just seems

third ph—l—ngesD —lthough thinD —llow in™re—sing the th—t the proportions of the front lim˜ of the wor—

˜e—ring surf—™eF sn the genus iquusD for ex—mpleD it g—vorso spe™imen —re the s—meD or very simil—rD —s

the ext—nt epennine ™h—moisF st ™—n ˜e —lso seen th—t h—s —lre—dy ˜een o˜served th—t the hemionesD the —sses

the r—tio r—diusGmet—™—rpus for the modern ‚F pyren—iE —nd the fur™hell9s ze˜r—sD whi™h live on h—rd groundsD

QWV ƒ—l—ri vFD ‚olfo wF pF 8 €etronio gF

MC PIP PIP PIP GPNP GPNP GPNP ANP ANP ANP parva parva parva max min mea max min mea max min mea max min mea Humerus / Metacarpus 1.261 1.308 1.275 1.292 1.271 1.228 1.251 0.991 0.946 0.969

Radius / Metacarpus 1.255 1.192 1.163 1.179 1.342 1.302 1.322 1.292 1.214 1.268 1.195 1.155 1.180

„—˜F V E ‚—tiosD —™™ording to g—p—sso f—r˜—to et —lF @IWVSAD ˜etween the gre—t lengths of front lim˜ ˜ones of the spe™imen from qrott— wor—

g—vorso ™omp—red with living ™h—mois —nd v—te €leisto™ene ™h—mois from €yrenees —nd s˜eri—n €eninsul—F wgX qrott— wor—

g—vorsoY €s€X €yrenees —nd s˜eri—n €eninsul—Y q€x€X ‚F rupi™—pr— rupi™—pr— from qr—n €—r—diso x—tion—l €—rkY ex€X ‚F pyren—i™—

orn—t— from e˜ruzzi x—tion—l €—rkY p—rv—X ‚F pyren—i™— p—rv— from the g—nt—˜ri—n r—ngeY m—xX m—ximumY minX minimumY me—X

me—nF

™— p—rv— is very simil—rD or the s—meD to th—t of the v—te ferred to ‚upi™—pr— spFD ™—n ˜e —ttri˜uted to ‚F pyren—iE

€leisto™ene ™h—mois from €yrenees —nd s˜eri—n €eninE ™— orn—t—F

sul—F gomp—risons with s—mples of ext—nt elpineD

epennine —nd €yrene—n ™h—mois —nd with fossil reE

m—ins from m—ny v—te €leisto™ene —nd older rolo™ene

gon™lusions

sites of st—ly —nd €yrenees —nd s˜eri—n €eninsul— h—ve

shown th—t the ˜ones of the spe™imen from wor— g—E

„he system—ti™s of the genus ‚upi™—pr— is ˜—sed

vorso @—s well —s most other v—te €leisto™ene spe™imensA on some fe—tures of the skull —nd of the hornE™ores @see

—re usu—lly gre—ter th—n those of rolo™ene —nd living ––„—xonomy of ‚upi™—pr— spe™ies99AF sn this p—perD —lso

™h—moisF roweverD we ™—n point out some pe™uli—ritiesF —mong the lim˜ ˜ones some morphometri™ differen™es

„he third ph—l—nges —nd the posterior first ph—E ˜etween the modern elpineD €yrene—n —nd epennine

l—nges —re rel—tively more elong—ted th—n to the ext—nt ™h—mois were notedF

™h—mois —nd fossil s—mples for ™omp—risonF „his p—rtiE „he skull —nd hornE™ore fe—tures of ™h—mois from

™ul—r fe—ture m—y ˜e —n —d—pt—tion to w—lk —nd run most qrott— wor— g—vorso —re typi™—l of the living epennine

™h—moisF elso the morphometry of the epiphyses of the of the ye—r on rel—tively soft —nd snowy soilsF purtherE

met—™—rpus —re simil—r to those of ext—nt epennine ™h—E moreD humerus —nd dist—l ti˜i— —re morphometri™—lly

moisD —lthough the morphologi™—l differen™es —lso in ™loser to those of the modern elpine ™h—mois th—n to

this ˜one ˜etween epennineD elpine —nd €yrene—n ™h—E of the epennine ™h—moisF w—y˜e this differen™e ™ould

mois —re not very noti™e—˜leF eddition—llyD the proporE ˜e due to norm—l intr—spe™ifi™ v—ri—˜ilityD ˜ut ™—nnot ˜e

tions of the front lim˜ h—ve v—lues ™loser to those of the ex™luded th—t the humerusD slightly ™ompressed in the

ext—nt epennine ™h—mois th—n to the elpine —nd €yrE —nteroEposterior dire™tion —nd less in the medioEl—ter—l

ene—n ™h—moisF por —ll these re—sons the su˜EentireD p—rE dire™tionD —nd the dist—l epiphysis of the ti˜i—D r—ther

ti—lly —rti™ul—ted skeleton from qrott— wor— g—vorsoD

squ—tD —re re™ent —d—pt—tions of modern epennine ™h—E

—n over IPEye—rs ™h—moisD for the l—rge size of its hornE

moisF sn su™h ™—seD we ™—n remem˜er th—t even the

™ore pro˜—˜ly — m—leD ™—n ˜e —s™ri˜ed to ‚upi™—pr— pyE

sm—ller ˜ody size of g—nt—˜ri—n ™h—mois —nd the perE

ren—i™— orn—t—F

pendi™ul—r insertion of hornE™ores of the elpine ™h—E

„he spe™imen from wor— g—vorso is to ˜e ™orreE

mois —re some fe—tures th—t would h—ve ˜een —™quired

l—ted to — temper—te os™ill—tion of wsƒ QD thus it is the

only during the rolo™ene @eltun— IWUPY w—sini IWVSAF first —s™ert—ined o™™urren™e of orn—t—Elike ™h—moisD so

f—r known with ™ert—inty only in the rolo™ene @‚ip— e™knowledgementsF ‡e wish to th—nk —ll the people who m—de

qr—nde —nd f—lzeD see w—sini IWVSY w—sini 8 vov—ri this p—per possi˜leD —llowing the study of the ™olle™tions —nd the reE

m—ins of ™h—mois stored in their snstitutionsF e spe™i—l th—nks to w—r— IWVVY voreti 8 ƒ—lerno IWVWA —nd possi˜ly in the v—teE

voreti of the qruppo ƒpeleologi™o of qu—ldo „—dinoD €—ol— „ollis of gl—™i—l @the two hornE™oresD one juvenileD without fronE

the inte eutonomo €—r™o x—zion—le d9e˜ruzzoD v—zio e wolise of

t—l ˜ones from ‚omito di €—p—sideroD ƒouthern epenE

€es™—sseroli —nd „onino „—gli—™ozzo of the wuseo x—zion—le €reistorE

ninesD northern g—l—˜ri—D see w—sini IWVSY w—sini 8

i™o itnogr—fi™o ––vuigi €igorini99 of ‚ome for their kindnessF ‡e —lso

vov—ri IWVVAF wost likely —lso the others ™h—mois reE th—nk —n —nonymous reviewer —nd pederi™o w—siniD for the useful

m—ins from ƒ IHSD ƒ IHR —nd ipigr—vetti—n l—yer of suggestions th—t improved the qu—lity of the m—nus™riptD —nd vetizi—

ƒilvestri of hurh—m niversityD for the review of our ˜roken inglishF qrott— wor— g—vorsoD prelimin—ry —nd ™—utiously reE

v—te €leisto™ene epennine ™h—mois QWW

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efeÂrez helg—do pFD wolero qutieÂrrez qF 8 fustos €retel †F ™i—le e del primo postgl—™i—leF snX ‡F‡FpF @idFA E n—

†it— per l— x—tur—X IHIEIPHD g—merino @wgAF @IWVIA E vos restos fo siles m—Âs meridion—les de ‚upiE

fenini eFD fos™—to €F 8 q—m˜—ssini €F @IWWUA E qrott— dell— ™—pr— rupi™—pr— h—ll—dos en iurop—F goloquios €—E

g—l— @ƒ—lernoAX industrie liti™he e f—une uluzzi—ne ed leontolFD QTX SQESWF

—urign—zi—neF ‚ivF ƒ™iF €reistFD RUX QUEWSF eltun— tF @IWUPA E p—un— de w—mõÂferos de los y—™imientos

fertolini wFD qurioli pFD ‚om—ndini wF 8 ƒ—l— fF @PHIPA E prehisto ri™os de quipu z™o—F wuni˜eD PRX IERTRF

xuove ™onsider—zioni p—leontologi™he e —r™heozooE eltun— tF @IWVIA E ‚estos oseos de y—™imiento prehistori™o

logi™he sui resti f—unisti™i del livello ipigr—vetti—no del ‚—s™—noF wonogrF gentro snvestF wusF elt—mir—D Ä

del ‚ip—ro di fi—rzo @ hAF snX he qrossi w—zzorin QX PPQEPTWF

tFD ƒ—™™—Á hF 8 „ozzi gF @idsA E etti del Të gonvegno eltun— tF @IWVTA E „he m—mm—li—n f—un—s from the prehisE

x—zion—le di er™heozoologi—D €—r™o dell9yre™™hiell— tori™ site of v— ‚ier—F snX ƒtr—us vFqF 8 gl—rk qF

ƒ—n ‚om—no in q—rf—gn—n— E vu™™—D PIEPR m—ggio @idsA E v— ‚ier— g—veD ƒtone ege runter q—therer

PHHWX PSEQPD vu™™—F ed—pt—tions in xorthern ƒp—inX PQUERWUD erizon—

fl—hout wF @IWUPA E ur „—xonomie der €opul—tion von ƒt—te niversity €ressF

‚upi™—pr— rupi™—pr— @vinneÂD IUSVA in der rohen „—E eltun— tF 8 w—riezkurren— uF @IWVRA E f—ses de su˜sisten™i—D

tr—F oolF listyD PX IISEIQPF

de origen —nim—lD de los po˜l—dores de ik—inF snX elE

fos™—to €F @IWWRA E qrott— €—gli™™iX l— f—un— — gr—ndi m—mE

tun— tF 8 werino tFwF @idsA E il y—™imiento preistorE

miferi degli str—ti PPEPR @qr—vetti—no —nti™o E euriE

i™o de l— guev— de ik—in @he˜—D quipu z™o—AX PIIEPVHD

gn—zi—noAF ‚ivF ƒ™iF €reistFD RTX IRSEIUTF

ƒo™ied—d de istudios †—s™osD quipuÂz™o—F

fos™—to €FD ‚on™hitelli eF 8 ‡ierer F @IWWUA E sl qr—vetE eltun— tF 8 w—riezkurren— uF @IWVSA E f—ses de su˜sisten™i—

ti—no —nti™o dell— qrott— dell— g—l— — w—rin— di g—E

de los po˜l—dores de irr—ll—X w—™rom—mõÂferosF wuE

merot—F €—letnologi— e —m˜ienteF ‚ivF ƒ™iF €reistFD RVX

ni˜eD QUX VUEIIUF

WUEIVTF eltun— tF 8 w—riezkurren— uF @PHHHA E w—™rom—mõÂferos del

fos™—to €F 8 ƒ—l— fF @IWVHA E h—ti p—leontologi™iD p—leoe™oE

y—™imiento de v—˜eko uo˜— @err—s—teD €—õÂs †—s™oAF

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g—mer—no vF @IWIR—A E ‚i™er™he intorno —i g—mos™iF g—E

mõÂferos del y—™imiento de —toy— @x—v—rr—AF „r—˜F

mos™io delle elpiD p—rte sF wemF ‚F e™™—dF ƒ™iF „orino

erqueolF x—v—rr—D ISX IHIEIIQF

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eltun— tFD w—riezkurren— uF 8 ilorz— wF @PHHPA E erqueoE

g—mer—no vF @IWIR˜A E ‚i™er™he intorno —i g—mos™iF g—E

zoologõ— de los nivelos p—leolõÂti™o de l— guev— de

mos™io delle elpiD p—rte ssF wemF ‚F e™™—dF ƒ™iF „orino

e˜—untz @err—izD x—v—rr—AF ƒ—ldvieD PX IEPTF

@glF ƒ™iF pisFD w—tFD x—tFAD TR@IRAX IEVVF

elv—rez „F FD wor—les eF 8 ƒese gF @IWWPA E w—miferos del

g—mer—no vF @IWIT—A E ‚i™er™he intorno —i g—mos™iF €—rte

y—™imiento del €leisto™eno ƒuperior de guev— will—n

sssF wemF ‚F e™™—dF ƒ™iF „orino @glF ƒ™iF pisFD w—tFD

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g—mer—no vF @IWIT˜A E s ™—r—tteri del ™r—nioD dell— ™olor—E

st—lyX — revised ™he™klist of spe™ies —nd su˜spe™ies

zione e delle ™orn— nell— distinzione dei g—mos™i in

@w—mm—li—AF ƒen™ken˜ergF ˜iolFD UWX PUIEPVTF

spe™ie e sottospe™ieF ‚ivF entropolF E etti ƒo™F ‚om—n—

entonioli pF 8 †—i qFfF @PHHRA E glimex m—ps st—lyD expl—E

entropolFD PH@SAX IEIRF

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RHR ƒ—l—ri vFD ‚olfo wF pF 8 €etronio gF Specimen n. Measurements (mm) according von den Driesch (1976) eppendix I

GL GB Bfcr Bfcd GLF H we—surements @mmA of postE™r—E

Atlas 2289 50.7 43.2 41.5 42.3 31.5 ni—l rem—ins of epennine ™h—mois from

LCDe LAPa Bfcr BPacd BPtr SBV Bfcd H qrott— wor— g—vorsoF

Axis 2731 55.4 49.5 39.4 27.4 24.2 22.0 45.5 we—surement —˜˜revi—tionsX etl—sX

HS DHA Ld SLC GLP LG BG qvX gre—test lengthY qfX gre—test ˜re—dth

Scapula 2285 29.1 26.3 21.1 over the wingsY fp™rX ˜re—dth of ™r—ni—l

GL GLC Bp Dp SD Bd Dd BT —rti™ul—r surf—™eY fp™dX ˜re—dth of ™—ud—l Humerus 2204 193.0 178.0 46.0 13.2 31.4 29.3 30.0 —rti™ul—r surf—™eY qvpX gre—test length

Humerus 2208 193.0 178.0 40.0 45.6 13.6 32.2 29.3 30.3 from the f—™ies —rti™ul—ris ™r—ni—lis to the

Femur 2156 214.0 47.0 21.2 16.1 40.1 48.9 ™—ud—lisY rX heightY exisX vgheX length in

the region of the ˜ody in™luding the densY GL LO DPA SDO BPC ve€—X gre—test length of the —r™Y fp™rX

Ulna 2191 37.3 28.4 23.6 14.3 ˜re—dth of ™r—ni—l —rti™ul—r surf—™eY f€—™dX

GL Bp Dp SD Bd Dd ˜re—dth —™ross the pro™essus —rti™ul—res

Radius 2261 192.0 32.5 18.2 17.9 29.4 20.4 ™—ud—lesY f€trX ˜re—dth —™rossthe tr—nsE

Radius 2191a 17.8 29.4 20.2 verse pro™essusY ƒf†X sm—llest ˜re—dthY

Tibia 2197 15.1 28.6 19.1 fp™dX ˜re—dth of ™—ud—l —rti™ul—r surf—™eY

Tibia 2199 28.6 19.3 rX heightY ƒ™—pul—X rX height —long the Metatacarpus 2249 153.0 23.9 18.5 16.1 29.1 17.3 spineY hreX di—gon—l heightY vdX dors—l

Metatacarpus 2232 24.1 18.7 lengthY ƒvgX sm—llest length of ™ollum s™—E

Metatacarpus 2258 29.2 17.3 pul—eY qv€X gre—test length of glenoid proE ™essY vqX length of glenoid ™—vityY fqX

Metatarsus 2196 22.7 20.8 14.9 ˜re—dth of glenoid ™—vityY rumerus —nd

Metatarsus 2201 22.8 20.8 15.0 pemurX qvX gre—test lengthY qvgX gre—test

GLI GLm Dl Dm Bd length from ™—putY fpX ˜re—dth of proxim—l

Talus 2264 31.4 30.5 17.7 18.2 20.5 endY hpX depth of the proxim—l end @in the

Talus 2218a 31.1 30.2 17.8 18.2 20.5 femur only ™—putAY ƒhX sm—llest ˜re—dth of

Talus * 2315 27.5 26.8 14.9 15.7 18.4 the di—physisY fdX ˜re—dth of dist—l endY

GL GB hdX depth of the dist—l endY f„X ˜re—dth Calcaneus 2255 65.1 22.2 of the tro™le— @only humerusAY ln—X qvX

Calcaneus 2265 66.6 22.3 gre—test lengthY vyX length of the ole™r—E

Centrotarsale 2256 24.4 nonY h€eX depth —™ross the pro™essus —nE ™on—eusY ƒhyX sm—llest depth of ole™r—E

GL Bp SD Bd nonY f€gX ˜re—dth —™ross the ™oronoid

Phalanx Ia 2207 47.6 14.3 9.0 13.4 pro™essY ‚—diusD „i˜i—D wet—™—rpusD wet—E

Phalanx Ia 2260 47.3 13.3 10.5 13.1 t—rsusX qvX gre—test lengthY fpX ˜re—dth of

Phalanx Ia 2284 47.1 13.4 10.5 13.1 proxim—l endY hpX depth of the proxim—l

Phalanx Ia 2291 46.3 13.6 11.2 13.3 endY ƒhX sm—llest ˜re—dth of the di—physisY

Phalanx Ip 2202 52.4 13.8 10.0 12.9 fdX ˜re—dth of dist—l endY hdX depth of the Phalanx Ip 2251 52.3 13.7 9.9 13.4 dist—l endY „—lusX qvlX gre—test length of

Phalanx Ip * 41 46.4 11.6 8.2 11.4 l—ter—l h—lfY qvmX gre—test length of medi—l

Phalanx II 2198 30.6 12.6 9.0 10.9 h—lfY hlX depth of l—ter—l h—lfY hmX depth of

Phalanx II 2202 a 32.7 13.3 9.0 9.9 medi—l h—lfY fdX ˜re—dth of dist—l endY g—lE ™—neus —nd ™entr—l t—rs—l ˜oneX qvX gre—tE

Phalanx II 2202 b 32.6 13.6 8.8 10.1 est length @only ™—l™—neusAY qfX gre—test

Phalanx II 2218b 31.1 13.2 9.1 10.8 ˜re—dthY pirst —nd ƒe™ond €h—l—nx @—X —nteE

Phalanx II 2251a 32.6 13.6 8.7 10.1 riorY pX posteriorAX qvX gre—test lengthY fpX

Phalanx II 2258a 30.5 13.3 8.7 10.8 ˜re—dth of proxim—l endY ƒhX sm—llest

DLS Ld MBS ˜re—dth of the di—physisY fdX ˜re—dth of

Phalanx III 2218c 40.6 31.2 8.3 dist—l endY „hird €h—l—nxX hvƒX di—gon—l Phalanx III 2258b 41.7 31.3 8.5 length of the soleY vdX length of dors—l surE

Phalanx III 2240 42.2 7.9 f—™eY wfƒX ˜re—dth in the middle of the

Phalanx III 2251b 8.2 soleF B from IHR ƒ D this spe™imens —re not of the su˜Eentire skeletonF

v—te €leisto™ene epennine ™h—mois RHS eppendix P site Specimen n. Measurements (mm) according von den Driesch (1976)

GL GB Bfcr Bfcd GLF H we—surements @mmA t—ken diE

re™tly ˜y the —uthors on postE™r—ni—l Ba Atlas wn 40.0 42.0 29.5

˜ones of the ™h—mois rem—ins of v—te SM Atlas P136/5 47.4 60.7 39.6 37.4 43.2 34.0

€leisto™ene —nd e—rly rolo™ene from LCDe LAPa Bfcr BPacd BPtr SBV Bfcd H

qrott— delle f—lze @f—AD housed —t the SM Axis P136/6 37.2

qu—ldo „—dino ƒpeleologi™—l qroupD snE HS DHA Ld SLC GLP LG BG

g—r—no @snAD housed in the i—rth ƒ™ien™e SM Scapula P136/7 22.7 34.9 28.3 23.1 hep—rtment of ––ƒ—pienz—99 niversity of GL GLC Bp Dp SD Bd Dd BT

‚omeD ‚ip—ro di pum—ne @puAD ‚ip—ro ƒoE Ba Humerus wn 175.0 165.0 45.0 15.2 25.0

m—n @ƒoA —nd ƒ—ntu—rio dell— w—donn— Ba Humerus wn 15.5 32.0 23.6 30.5

@ƒwAD stored in the x—tion—l wuseum of Ba Humerus wn 15.4 31.8 24.2 29.0 €rehistory —nd ithnogr—phy ––vuigi €iE

gorini99 of ‚ome @for —˜˜revi—tions of In Humerus FG45 28.1 25.7 26.4 me—surements see ™—ption of eppendix IAF SM Humerus P136/8 33.3 29.0 31.0 SM Humerus P136/9 32.8 27.4 30.6 GL LO DPA SDO BPC Ba Ulna wn 25.0 21.5 14.8 SM Ulna P136/10 43.6 28.2 24.2 18.0 GL Bp Dp SD Bd Dd Ba Radius wn 170.0 28.8 18.0 17.0 28.6 21.0 Ba Tibia wn 15.7 21.5 18.2 So Tibia A51-20 32.4 21.8 Ba Metatacarpus wn 15.7 27.7 16.4 SM Metatacarpus P136/12 29.4 17.5 Ba Metatarsus wn 153.0 24.2 23.6 15.8 28.3 18.9 Ba Metatarsus wn 155.0 21.2 21.0 14.3 26.5 17.4 SM Metatarsus P136/15 29.7 19.8 GLI GLm Dl Dm Bd Ba Talus wn 30.8 30.0 17.7 18.3 20.0 SM Talus P136/18 33.0 31.5 17.9 19.8 20.7 So Talus RS360 34.3 33.0 19.2 21.1 22.0 GL GB Ba Calcaneus wn 20.1 In Calcaneus FG43 63.5 22.5 SM Calcaneus P136/16 70.5 24.0 SM Calcaneus P136/17 63.2 22.4 Ba Centrotarsale wn 22.5 SM Centrotarsale P136/19 26.4 SM Centrotarsale P136/20 25.2 GL Bp SD Bd Ba Phalanx I wn 40.7 12.6 10.7 12.2 Ba Phalanx I wn 45.1 12.7 10.5 12.3 Ba Phalanx I wn 45.1 12.6 10.1 12.2 SM Phalanx I P136/22 13.6 8.9 SM Phalanx I P136/23 13.1 SM Phalanx I P136/25 14.2 Ba Phalanx II wn 28.3 12.0 8.8 10.8 SM Phalanx II P136/27 31.3 12.7 8.3 10.2 SM Phalanx II P136/28 31.6 14.2 8.9 10.6 DLS Ld MBS Fu Phalanx III P6316 34.7 28.8 7.8 SM Phalanx III P136/29 33.4 28.7 7.4 SM Phalanx III P136/30 33.8 27.9 7.3

RHT ƒ—l—ri vFD ‚olfo wF pF 8 €etronio gF site Specimen n. Measurements (mm) according von den Driesch (1976) eppendix Q GL GB Bfcr Bfcd GLF H

ANP Atlas we—surements @mmA t—ken diE 27 47.3 57.6 40.5 40.0 41.2 29.4 re™tly ˜y the —uthors on postE™r—ni—l

ANP Atlas 35 42.3 52.5 37.2 36.7 38.4 26.7 ˜ones of epennine ™h—mois from e˜ruzzi

ANP Atlas 201 46.2 57.0 40.2 40.2 43.0 29.0 x—tion—l €—rk @ex€A housed in the ƒ™ienE

ANP Atlas 238 47.5 59.2 41.4 40.0 41.5 30.5 tifi™ ƒervi™e of ––inte eutonomo €—r™o

ANP Atlas wn 45.2 52.6 39.0 39.4 45.5 30.5 x—zion—le d9e˜ruzzoD v—zio e wolise99

GPNP Atlas wn 45.7 56.8 41.4 38.3 39.6 31.1 of €es™—sseroli @v9equil—A —nd of elpine LCDe LAPa Bfcr BPacd BPtr SBV Bfcd H ™h—mois from qr—n €—r—diso x—tion—l

ANP Axis 27 55.1 50.4 37.2 28.2 37.3 22.6 18.1 45.5 €—rk @q€x€A stored in the x—tion—l wuE

ANP Axis * 35 49.8 47.4 35.2 27.0 34.2 25.0 16.7 37.5 seum of €rehistory —nd ithnogr—phy

ANP Axis ––vuigi €igorini99 of ‚ome @for —˜˜revi—E 58 54.2 46.2 38.6 30.0 38.5 28.2 18.8 41.3 tions of me—surements see ™—ption of epE

ANP Axis 201 54.2 49.5 38.0 29.0 37.0 21.8 18.5 45.0 pendix IAF Bspe™imens with the proxim—l

ANP Axis 238 56.5 50.8 37.8 28.5 38.0 22.2 18.8 41.5 —ndGor dist—l epiphysis not ™ompletely

ANP Axis 239 52.7 46.1 34.9 23.8 34.2 20.8 16.4 42.2 fused —ndD thereforeD ex™luded ˜y st—tistiE ANP Axis * wn 54.0 37.5 25.0 35.5 22.5 17.5 41.5 ™—l —n—lysesF GPNP Axis wn 52.4 36.1 24.6 31.8 18.5 19.7 42.3 HS DHA Ld SLC GLP LG BG ANP Scapula 26 154.0 165.0 92.5 21.1 30.3 27.7 22.7 ANP Scapula 35 139.0 150.0 85.0 18.8 30.2 24.4 20.0 ANP Scapula 47 158.0 170.0 96.0 22.0 30.5 27.0 22.7 ANP Scapula 201 151.0 164.0 93.5 19.0 31.5 25.0 21.0 ANP Scapula 238 158.0 170.0 97.5 21.5 31.4 26.1 22.5 ANP Scapula 239 147.0 158.0 84.5 18.8 30.7 26.8 20.6 ANP Scapula wn 144.0 156.0 85.0 19.0 30.5 24.2 21.6 GPNP Scapula P506/73 157.0 171.0 98.0 20.5 32.5 26.6 22.3 GPNP Scapula wn 147.0 163.0 93.0 20.2 28.1 25.5 20.9 GL GLC Bp Dp SD Bd Dd BT ANP Humerus 26 180.5 165.0 38.4 49.8 14.8 34.4 26.7 29.7 ANP Humerus * 35 169.0 153.0 35.0 41.6 13.3 31.2 25.4 28.0 ANP Humerus 47 178.0 163.0 38.7 44.0 14.4 34.6 27.0 31.3 ANP Humerus 191 178.0 164.0 43.6 14.8 37.7 25.6 29.7 ANP Humerus 201 179.0 36.5 43.5 14.8 32.2 25.5 29.5 ANP Humerus 238 181.0 176.0 38.2 43.5 16.0 34.5 27.8 31.2 ANP Humerus 239 170.0 157.0 37.2 43.1 14.4 33.3 25.1 28.2 ANP Humerus * wn 171.0 157.0 36.8 43.2 13.8 32.2 25.5 30.5 GPNP Humerus P506/75 190.0 176.0 40.0 46.5 13.5 32.5 29.5 30.5 GPNP Humerus wn 191.0 173.0 38.8 46.2 13.8 31.8 26.8 29.7 ANP Femur 21 210.0 20.5 17.5 42.2 49.4 ANP Femur 26 206.0 20.5 15.9 38.2 47.8 ANP Femur * 35 199.0 42.0 19.0 16.0 38.8 43.2 ANP Femur 58 211.5 210.0 45.7 21.7 17.1 38.8 47.6 ANP Femur 201 205.0 200.0 44.4 20.2 18.2 39.1 47.8 ANP Femur 238 220.0 214.0 47.2 21.1 16.8 40.4 49.9 ANP Femur 239 196.0 188.0 43.6 20.0 16.5 38.2 46.8 ANP Femur 1100 203.0 44.0 20.4 16.8 38.3 47.8 ANP Femur * wn 205.0 200.0 45.2 20.3 17.5 39.6 45.0 GPNP Femur P506/120 216.0 212.0 49.4 21.7 16.5 41.9 49.7 GPNP Femur wn 215.0 209.0 45.0 20.2 16.8 40.3 47.2 GL LO DPA SDO BPC ANP Ulna 26 228.0 36.1 22.2 24.2 15.6 ANP Ulna * 35 34.6 22.3 20.0 15.0 ANP Ulna 47 36.7 24.8 22.2 15.3 ANP Ulna 191 224.0 34.8 22.3 24.6 16.8 ANP Ulna 201 224.0 32.1 21.5 24.7 15.6 ANP Ulna 238 221.0 32.0 23.4 26.4 15.5 ANP Ulna 239 210.0 34.5 21.1 23.9 14.8 ANP Ulna * wn 220.0 37.5 20.8 22.8 14.3

v—te €leisto™ene epennine ™h—mois RHU

site Specimen n. Measurements (mm) according von den Driesch (1976) GPNP Ulna P506/79 36.6 25.6 24.5 17.8 GL Bp Dp SD Bd Dd ANP Radius 26 178.0 29.4 17.4 17.3 28.8 20.4 ANP Radius * 35 160.0 27.5 15.2 15.2 25.2 20.4 ANP Radius 47 176.0 33.6 16.5 17.5 28.4 19.7 ANP Radius 191 183.0 29.2 17.4 17.4 27.6 19.8 ANP Radius 201 184.0 28.8 16.2 17.5 28.6 19.4 ANP Radius 238 186.0 31.2 17.1 17.4 28.4 20.2 ANP Radius 239 174.0 28.5 16.1 16.5 27.2 18.9 ANP Radius * wn 182.0 28.8 17.5 17.8 27.5 21.6 GPNP Radius P506/77 194.0 30.5 17.0 17.8 29.9 22.4 GPNP Radius wn 196.0 29.2 15.9 16.5 29.4 19.5 ANP Tibia 21 272.0 43.3 44.2 14.7 26.7 21.2 ANP Tibia 26 266.0 43.1 42.2 16.0 27.2 20.2 ANP Tibia 33 253.0 41.4 40.2 19.2 26.5 20.6 ANP Tibia * 35 252.0 39.8 39.0 15.6 24.2 20.0 ANP Tibia 58 272.0 42.2 43.6 16.0 28.2 20.8 ANP Tibia 201 265.0 43.4 42.6 15.5 26.6 20.4 ANP Tibia 238 268.0 43.6 44.4 15.6 27.0 22.4 ANP Tibia 239 254.0 42.5 41.1 15.2 25.5 18.4 ANP Tibia * wn 265.0 43.2 42.3 16.0 26.5 20.8 GPNP Tibia P506/124 273.0 47.2 41.0 15.5 28.8 19.1 GPNP Tibia wn 273.0 43.0 42.0 15.8 27.5 19.8 ANP Metacarpus 26 142.0 24.5 17.3 15.8 29.2 17.0 ANP Metacarpus 26 143.0 24.5 17.1 15.7 29.4 16.8 ANP Metacarpus * 35 139.0 22.8 16.2 15.3 29.0 15.8 ANP Metacarpus * 35 139.0 23.4 16.2 15.0 28.3 15.4 ANP Metacarpus 47 145.0 24.8 17.7 15.0 30.0 16.6 ANP Metacarpus 191 143.0 24.7 16.9 16.0 29.4 16.9 ANP Metacarpus 191 143.0 24.5 16.8 16.0 29.2 16.9 ANP Metacarpus 201 143.0 24.0 17.0 16.0 28.8 16.8 ANP Metacarpus 201 143.0 24.2 17.2 16.0 28.8 16.8 ANP Metacarpus 238 144.0 25.5 17.7 15.8 30.5 17.2 ANP Metacarpus 239 135.5 23.0 16.5 15.2 28.0 16.0 ANP Metacarpus * wn 142.0 24.2 17.0 16.8 28.2 16.5 GPNP Metacarpus P506/93 149.0 24.5 18.3 15.3 29.0 17.8 GPNP Metacarpus wn 146.0 24.1 18.4 14.6 27.9 17.8 ANP Metatarsus 21 165.0 24.2 22.2 14.3 29.2 19.3 ANP Metatarsus 26 162.0 22.8 19.3 13.8 28.5 17.8 ANP Metatarsus * 35 157.5 21.8 18.8 14.2 21.7 17.3 ANP Metatarsus 58 163.0 22.4 20.2 14.6 29.3 18.9 ANP Metatarsus 201 163.0 22.7 21.1 14.4 27.9 15.4 ANP Metatarsus 238 161.0 23.0 20.0 14.2 28.8 19.2 ANP Metatarsus 239 153.0 21.1 20.5 13.5 26.5 16.9 ANP Metatarsus * wn 160.0 22.8 19.5 14.5 27.2 18.6 GPNP Metatarsus P506/138 171.0 22.1 20.6 13.8 28.9 18.5 GPNP Metatarsus wn 179.0 22.0 20.1 13.4 26.6 17.8 GLI GLm Dl Dm Bd ANP Talus 21 29.2 29.0 16.2 18.0 20.6 ANP Talus 26 28.5 28.2 16.3 17.6 19.9 ANP Talus 35 27.7 27.4 15.8 16.2 19.4 ANP Talus 58 28.5 28.1 17.4 18.3 21.2 ANP Talus 201 28.1 27.6 17.5 18.1 20.5 ANP Talus 238 30.1 29.2 17.3 18.6 20.0 ANP Talus 239 27.5 27.3 15.8 17.0 19.2 ANP Talus wn 28.2 28.0 16.5 17.0 19.5 GPNP Talus P506/132 31.2 30.3 17.5 18.5 19.8 GPNP Talus wn 29.8 29.2 17.4 17.8 19.2 GL GB ANP Calcaneus 21 63.0 23.5

RHV ƒ—l—ri vFD ‚olfo wF pF 8 €etronio gF

site Specimen n. Measurements (mm) according von den Driesch (1976) ANP Calcaneus 26 60.6 22.0 ANP Calcaneus 35 58.2 19.6 ANP Calcaneus 58 61.5 21.2 ANP Calcaneus 201 61.5 20.2 ANP Calcaneus 238 62.6 22.2 ANP Calcaneus 239 59.4 21.5 ANP Calcaneus * wn 59.0 20.7 GPNP Calcaneus P506/133 65.6 21.5 ANP Centrotarsale 21 26.4 ANP Centrotarsale 35 24.1 ANP Centrotarsale 58 25.4 ANP Centrotarsale 201 24.5 ANP Centrotarsale 238 25.4 ANP Centrotarsale wn 24.8 GPNP Centrotarsale P506/134 25.2 GPNP Centrotarsale wn 23.5 GL Bp SD Bd ANP Phalanx Ia 26 42.5 12.8 11.1 13.2 ANP Phalanx Ia 26 42.7 13.4 10.7 13.2 ANP Phalanx Ia 35 41.2 13.4 9.9 12.4 ANP Phalanx Ia 201 43.1 12.9 11.1 12.8 ANP Phalanx Ia 238 45.0 14.6 11.2 13.5 ANP Phalanx Ia 239 42.0 12.5 10.2 12.0 ANP Phalanx Ia wn 41.8 13.5 10.5 12.1 GPNP Phalanx Ia P506/101 45.8 13.3 10.0 12.7 GPNP Phalanx Ia wn 41.5 13.4 9.9 11.8 ANP Phalanx Ip 21 47.0 13.5 10.0 12.0 ANP Phalanx Ip 26 47.0 12.8 10.8 12.5 ANP Phalanx Ip 35 42.7 12.4 9.3 11.7 ANP Phalanx Ip 58 47.0 13.0 10.2 12.7 ANP Phalanx Ip 201 46.5 12.5 10.7 12.4 ANP Phalanx Ip 238 46.2 13.4 10.5 12.0 ANP Phalanx Ip 239 45.5 12.5 9.5 12.0 ANP Phalanx Ip wn 44.2 13.1 9.2 11.6 GPNP Phalanx Ip P506/146 49.1 13.4 9.3 12.2 GPNP Phalanx Ip wn 46.0 13.4 9.1 11.7 ANP Phalanx II 21 31.8 13.0 8.6 10.0 ANP Phalanx II 26 28.8 13.5 9.4 10.3 ANP Phalanx II 26 28.8 13.1 9.3 10.5 ANP Phalanx II 201 29.1 13.2 8.8 10.2 ANP Phalanx II 238 29.8 13.7 9.4 10.5 ANP Phalanx II 239 29.5 11.8 8.5 9.5 ANP Phalanx II wn 28.6 12.5 8.6 9.5 GPNP Phalanx II P506/103 30.7 12.0 8.8 10.1 GPNP Phalanx II P506/147 31.2 12.4 8.0 10.0 DLS Ld MBS ANP Phalanx III 26 34.1 25.3 8.2 ANP Phalanx III 26 33.5 24.3 7.9 ANP Phalanx III 26 32.8 25.8 7.2 ANP Phalanx III 26 32.7 25.3 7.2 ANP Phalanx III 201 35.2 26.5 7.4 ANP Phalanx III 238 33.3 25.3 7.6 ANP Phalanx III 239 34.0 25.0 7.5 ANP Phalanx III wn 32.2 24.2 6.9 GPNP Phalanx III P506/148 36.5 30.6 7.2